Minimal Effects of an Invasive Flowering Shrub on the Pollinator Community of Native Forbs

Biological invasions can strongly influence species interactions such as pollination. Most of the documented effects of exotic plant species on plant-pollinator interactions have been observational studies using single pairs of native and exotic plants, and have focused on dominant exotic plant species. We know little about how exotic plants alter interactions in entire communities of plants and pollinators, especially at low to medium invader densities. In this study, we began to address these gaps by experimentally removing the flowers of a showy invasive shrub, Rosa multiflora, and evaluating its effects on the frequency, richness, and composition of bee visitors to co-flowering native plants. We found that while R. multiflora increased plot-level richness of bee visitors to co-flowering native plant species at some sites, its presence had no significant effects on bee visitation rate, visitor richness, bee community composition, or abundance overall. In addition, we found that compared to co-flowering natives, R. multiflora was a generalist plant that primarily received visits from generalist bee species shared with native plant species. Our results suggest that exotic plants such as R. multiflora may facilitate native plant pollination in a community context by attracting a more diverse assemblage of pollinators, but have limited and idiosyncratic effects on the resident plant-pollinator network in general.     

Heterospecific pollen transfer from an exotic plant to native plants: assessing reproductive consequences in an Andean grassland

Background: The presence of exotic plants increases the heterospecific pollen (HP hereafter) received by native plants and reduces their reproductive output.

Aims: We assessed whether the exotic herb Echium vulgare (Boraginaceae) increased HP and reduced seed output of the native plants Phacelia secunda (Boraginaceae) and Stachys albicaulis (Lamiaceae) in an Andean locality, central Chile.

Methods: The presence of HP was studied in native plants growing with and without co-existence with the exotic E. vulgare. A complementary hand pollination experiment was carried out to assess whether E. vulgare pollen reduced the reproductive success of native plants.

Results: In the presence of E. vulgare, 17.3% and 3.7% of P. secunda and S. albicaulis individuals that coexisted with the exotic species received HP. For P. secunda, the number of conspecific pollen grains decreased in invaded patches compared with non-invaded patches; no differences were observed for S. albicaulis. The pollen of E. vulgare negatively affected the reproductive success of S. albicaulis but not that of P. secunda.

Conclusions: The presence of HP cannot be predicted from the presence of exotic plants alone, and other factors, such as flower morphology, could explain the greater HP transfer in P. secunda (actinomorphic flowers) than in S. albicaulis (zygomorphic flowers). A higher negative effect of E. vulgare pollen on P. secunda versus S. albicaulis could be related to the phylogenetic resemblance between the exotic donor and native recipient plant because pollen-stigma compatibility may be evolutionary conserved through common lineages.     

Effect of invader removal: pollinators stay but some native plants miss their new friend

Removal of invasive species often benefits biological diversity allowing ecosystems’ recovery. However, it is important to assess the functional roles that invaders may have established in their new areas to avoid unexpected results from species elimination. Invasive animal-pollinated plants may affect the plant–pollination interactions by changing pollinator availability and/or behaviour in the community. Thus, removal of an invasive plant may have important effects on pollinator community that may then be reflected positive or negatively on the reproductive success of native plants. The objective of this study was to assess the effect of removing Oxalis pes-caprae, an invasive weed widely spread in the Mediterranean basin, on plant–pollinator interactions and on the reproductive success of co-flowering native plants. For this, a disturbed area in central Portugal, where this species is highly abundant, was selected. Visitation rates, natural pollen loads, pollen tube growth and natural fruit set of native plants were compared in the presence of O. pes-caprae and after manual removal of their flowers. Our results showed a highly resilient pollination network but also revealed some facilitative effects of O. pes-caprae on the reproductive success of co-flowering native plants. Reproductive success of the native plants seems to depend not only on the number and diversity of floral visitors, but also on their efficiency as pollinators. The information provided on the effects of invasive species on the sexual reproductive success of natives is essential for adequate management of invaded areas.     

Increased relative abundance of an invasive competitor for pollination, Lythrum salicaria, reduces seed number in Mimulus ringens

When exotic plant species share pollinators with native species, competition for pollination may lower the reproductive success of natives by reducing the frequency and/or quality of visits they receive. Exotic species often become numerically dominant in plant communities, and the relative abundance of these potential competitors for pollination may be an important determinant of their effects on the pollination and reproductive success of co-occurring native species. Our study experimentally tests whether the presence and abundance of an invasive exotic, Lythrum salicaria L. (Lythraceae), influences reproductive success of a co-flowering native species, Mimulus ringens L. (Phrymaceae). We also examine the mechanisms of competition for pollination and how they may be altered by changes in competitor abundance. We found that the presence of Lythrum salicaria lowered mean seed number in Mimulus ringens fruits. This effect was most pronounced when the invasive competitor was highly abundant, decreasing the number of seeds per fruit by 40% in 2006 and 33% in 2007. Reductions in the number of seeds per fruit were likely due to reduced visit quality resulting from Mimulus pollen loss when bees foraged on neighboring Lythrum plants. This study suggests that visit quality to natives may be influenced by the presence and abundance of invasive flowering plants.     

Pollination of the European food-deceptive Traunsteinera globosa (Orchidaceae): the importance of nectar-producing neighbouring plants

European food-deceptive orchids generally flower early in spring and rely on naïve pollinators for their reproduction. Some species however, flower later in the summer, when many other rewarding plants species are also in bloom. In dense flowering communities, deceptive orchids may suffer from competition for pollinator resources, or might alternatively benefit from higher community attractiveness. We investigated the pollination strategy of the deceptive species Traunsteinera globosa, and more specifically whether it benefited from the presence of coflowering rewarding species. We carried out a population survey to quantify the density and reproductive success of the orchid as well as the density of all coflowering species. Our results suggest that the deceptive orchid not only benefited from the presence of coflowering species, but that interestingly the density of the species Trifolium pratense was significantly positively correlated with the orchid's reproductive success. This species might simply act as a magnet species attracting pollinators near T. globosa, or could influence the orchid reproductive fitness through a more species-specific interaction. We propose that morphological or colour similarities between the two species should be investigated in more detail to decipher this pollination facilitation effect.     

Comparing the reproductive success and pollination biology of an invasive plant to its rare and common native congeners: a case study in the genus <Emphasis Type="Italic">Cirsium</Emphasis> (Asteraceae)

Previous studies have examined an association between reproductive success and pollination biology of rare versus widespread species through pair-wise comparisons of native and invasive congeners or rare and common congeners. To determine the importance of reproductive success and pollination biology for an invasive thistle, Cirsium vulgare, we compared it in its invaded range to five, co-occurring native Cirsium species that range from rare to common. Native study species include C. fontinale var. fontinale, C. andrewsii, C. brevistylum, C. occidentale, and C. quercetorum. We compared all species’ reproductive success, insect visitation rate and composition, autonomous self-pollination, and level of pollen limitation in multiple populations. Species differed in their reproductive success; the invasive C. vulgare produced more flower heads per plant than most native species. C. vulgare attracted more visitors than its congeners. In addition, reproductive success and insect visitation significantly varied between populations within species, mainly due to aphid infestation in one population of C. occidentale. Unlike the rare species (C. fontinale and andrewsii), C. vulgare did not require a pollinator for high-levels of seed production. The remaining native species set fewer seeds than C. vulgare without a pollinator. However, differences in insect visitation and autonomous self-pollination did not lead to differences in pollen limitation across species or between populations. This result suggests that factors other than pollination biology determine the difference in reproductive success of these species. However, high levels of autonomous self-pollination and generalist insect visitation may allow the invasive C. vulgare to easily establish new populations from low numbers of propagules. Our study provides one contrast that should build towards a larger comparative analysis to examine general patterns in the relationship between reproductive success, pollination biology, rare and invasive species, and our ability to predict biological invasions in introduced species.     

Pollination in a patchily distributed lousewort is facilitated by presence of a co-flowering plant due to enhancement of quantity and quality of pollinator visits

Background and Aims

Plants surrounded by individuals of other co-flowering species may suffer a reproductive cost from interspecific pollen transfer (IPT). However, differences in floral architecture may reduce or eliminate IPT.

Methods

A study was made of Pedicularis densispica (lousewort) and its common co-flowering species, Astragalus pastorius, to compare reproductive and pollination success of lousewort plants from pure and mixed patches. Floral architecture and pollinator behaviour on flowers of the two plants were compared along with the composition of stigmatic pollen load of the louseworts. The extent of pollen limitation of plants from pure and mixed patches was also explored through supplemental pollination with self- and outcross pollen (PLs and PLx).

Key Results

Mixed patches attracted many more nectar-searching individuals of Bombus richardsi. These bumble-bees moved frequently between flowers of the two species. However, they pollinated P. densispica with their dorsum and A. pastorius with their abdomen. This difference in handling almost completely eliminated IPT. Lousewort plants from mixed patches yielded more seeds, and seeds of higher mass and germinability, than those from pure patches. Moreover, louseworts from mixed patches had lower PLs and PLx compared with those from pure patches.

Conclusions

Differences in floral architecture induced differences in pollinator behaviour that minimized IPT, such that co-flowering plants significantly enhanced quantity and quality of pollinator visits for the lousewort plants in patchy habitat. These findings add to our understanding of the mechanisms of pollination facilitation.     

Phylogenetic Community Context Influences Pollen Delivery to <Emphasis Type="Italic">Allium cernuum</Emphasis>

Few studies have examined how the number and identity of species in the neighbouring community influences the reproductive success of particular focal species. Pollen delivery, an important component of fitness of sexual plants, is a function of not just the floral traits of any particular individual, but of features of the population and community as it depends on pollinator abundance and preferences. Many pollinators in flowering communities will prefer patches with high floral abundance or diversity yet may exhibit lower floral constancy when more flowering species are present. Thus, pollination efficiency could increase or decrease with increased species richness and this will alter the selection pressures placed upon the floral traits (such as floral colour or reward) of any member of a particular community. Moreover, recent studies have indicated that plant-pollinator networks are phylogenetically structured (pollinators visit related plant species more than expected by chance) and this may be an important factor structuring flowering plant communities. Thus, the sheer number of species within a patch may be less important than the number of closely-related species. We investigate whether species richness or phylogenetic proximity of coflowering species influences the amount and proportion of conspecific pollen delivered to nodding onion, Allium cernuum, in fragment patches of Garry Oak meadows in South Western British Columbia, Canada. We find that pollen delivery depended upon the presence of close relatives far more than on species richness or population density, indicating a central role of the community structure on pollination in flowering plant communities. Insofar as pollen delivery relates to the relative number of seeds produced by members of the community, pollination may in turn determine the community structure of the next generation.     

Alien pollen grains interfere with the reproductive success of native congener

The effects of invasive species on native species comprise important conservation issues. Determining the mechanisms by which invasives exclude natives is indispensable to efficiently control their impact, but most invasives remain poorly studied. The purpose of this study was to elucidate potentially important but neglected mechanisms, reproductive interference, in wild Taraxacum systems, in which invasive Taraxacum officinale has displaced its native congener T. japonicum in Japan. Hand-pollination of mixed pollen grains significantly reduced the native seed-set compared to conspecific-only pollination. Moreover, natives with a high ratio of invasive pollen on their stigmas suffered severe seed-set reduction, and the proportion of invasive pollen on native stigmas increased as frequencies of the alien neighbor increased. These results, combined with those of previous studies, revealed that depositing invasive pollen on native stigmas contributes to the observed alien-frequency-dependent reduction of native seed-set, and strongly suggest that reproductive interference was the primary cause of displacement in the Taraxacum systems.     

Direct effects of a biocontrol agent are greater than indirect effects through flower visitors for the alien plant Dalmatian toadflax (Linaria dalmatica: Scrophulariaceae)

Herbivory and pollination are important determinants of female reproductive success in flowering plants. Plants must interact with herbivores and flower visitors simultaneously and interaction with one may alter the outcome of the interaction with the other. These indirect effects can have dramatic impacts on plant fitness. The purpose of this study was to examine whether the stem-boring weevil Mecinus janthiniformis (Curculionidae: Coleoptera) affects flower visitation rate and seed set of the exotic plant Dalmatian toadflax (Linaria dalmatica (L.) Mill. Scrophulariaceae). We compared the flower production, flower morphology, visitation rate, fruit production, and pollen limitation on Dalmatian toadflax plants with and without larval feeding by M. janthiniformis. Feeding by M. janthiniformis reduced the number of flowers and per plant visitation rate, and there was a significant interaction between herbivory and flower number suggesting that the change in visitation rate was not solely a function of a reduction in flower abundance. Herbivory also had direct negative impacts on the reproductive success of Dalmatian toadflax. Total flower and fruit production decreased by over 30 % in plants attacked by M. janthiniformis. However, plants with M. janthiniformis were not more pollen-limited than those without M. janthiniformis. This suggests that herbivory had primarily direct effects female reproductive success.     

The pollination efficiency of a pollinator depends on its foraging strategy,flowering phenology,and the flower characteristics of a plant species

Honey bees and stingless bees are generalist visitors of several wild and cultivated plants. They forage with a high degree of floral fidelity and thereby help in the pollination services of those plants. We hypothesized that pollination efficiency might be influenced by flowering phenology, floral characteristics, and resource collection modes of the worker bees. In this paper, we surveyed the foraging strategies of honey bees (Apis cerana, Apis dorsata, and Apis florea) and stingless bees (Tetragonula iridipennis) concerning their pollination efficiencies. Bees showed different resource gathering strategies, including legitimate (helping in pollination as mixed foragers and specialized foragers) and illegitimate (serving as nectar robbers and pollen thieves) types of flower visitation patterns. Foraging strategies are influenced by the shape of flowers, the timing of the visitation, floral richness, and bee species. Honey bees and stingless bees mainly acted as legitimate visitors in most plants studied. Sometimes honey bees served as nectar robbers in tubular flowers and stingless bees as pollen thieves in large-sized flowers. Among the legitimate categories, mixed foragers have a comparatively lower flower visitation rate than the specialized nectar and pollen foragers. However, mixed foragers have greater abundance and higher values of the single-visit pollination efficiency index (PEi) than nectar and pollen foragers. The value of the combined parameter ‘importance in pollination (PI)’ was thus higher in mixed foragers than in nectar and pollen foragers.     

Grassland invaders and their mycorrhizal symbionts: a study across climate and invasion gradients

Controlled experiments show that arbuscular mycorrhizal fungi (AMF) can increase competitiveness of exotic plants, potentially increasing invasion success. We surveyed AMF abundance and community composition in Centaurea stoebe and Potentilla recta invasions in the western USA to assess whether patterns were consistent with mycorrhizal-mediated invasions. We asked whether (1) AMF abundance and community composition differ between native and exotic forbs, (2) associations between native plants and AMF shift with invading exotic plants, and (3) AMF abundance and/or community composition differ in areas where exotic plants are highly invasive and in areas where they are not. We collected soil and roots from invaded and native forb communities along invasion gradients and in regions with different invasion densities. We used AMF root colonization as a measure of AMF abundance and characterized AMF communities in roots using 454-sequencing of the LSU-rDNA region. All plants were highly colonized (>60%), but exotic forbs tended to be more colonized than natives (P < 0.001). We identified 30 AMF operational taxonomic units (OTUs) across sites, and community composition was best predicted by abiotic factors (soil texture, pH). Two OTUs in the genera Glomus and Rhizophagus dominated in most communities, and their dominance increased with invasion density (r = 0.57, P = 0.010), while overall OTU richness decreased with invasion density (r = −0.61, P = 0.006). Samples along P. recta invasion gradients revealed small and reciprocal shifts in AMF communities with >45% fungal OTUs shared between neighboring native and P. recta plants. Overall, we observed significant, but modest, differences in AMF colonization and communities between co-occurring exotic and native forbs and among exotic forbs across regions that differ in invasion pressure. While experimental manipulations are required to assess functional consequences, the observed patterns are not consistent with those expected from strong mycorrhizal-mediated invasions.     

Interspecific pollinator movements reduce pollen deposition and seed production in Mimulus ringens (Phrymaceae)

Movement of pollinators between coflowering plant species may influence conspecific pollen deposition and seed set. Interspecific pollinator movements between native and showy invasive plants may be particularly detrimental to the pollination and reproductive success of native species. We explored the effects of invasive Lythrum salicaria on the reproductive success of Mimulus ringens, a wetland plant native to eastern North America. Pollinator flights between these species significantly reduced the amount of conspecific pollen deposited on Mimulus stigmas and the number of seeds in Mimulus fruits, suggesting that pollen loss is an important mechanism of competition for pollination. Although pollen loss is often attributed to pollen wastage on heterospecific floral structures, our novel findings suggest that grooming by bees as they forage on a competitor may also significantly reduce outcross pollen export and seed set in Mimulus ringens.     

Generalization versus Specialization in Pollination Systems: Visitors,Thieves, and Pollinators of Hypoestes aristata (Acanthaceae)

Many recent studies have suggested that the majority of animal-pollinated plants have a higher diversity of pollinators than that expected according to their pollination syndrome. This broad generalization, often based on pollination web data, has been challenged by the fact that some floral visitors recorded in pollination webs are ineffective pollinators. To contribute to this debate, and to obtain a contrast between visitors and pollinators, we studied insect and bird visitors to virgin flowers of Hypoestes aristata in the Bamenda Highlands, Cameroon. We observed the flowers and their visitors for 2-h periods and measured the seed production as a metric of reproductive success. We determined the effects of individual visitors using 2 statistical models, single-visit data that were gathered for more frequent visitor species, and frequency data. This approach enabled us to determine the positive as well as neutral or negative impact of visitors on H. aristata’s reproductive success. We found that (i) this plant is not generalized but rather specialized; although we recorded 15 morphotaxa of visitors, only 3 large bee species seemed to be important pollinators; (ii) the carpenter bee Xylocopa cf. inconstans was both the most frequent and the most effective pollinator; (iii) the honey bee Apis mellifera acted as a nectar thief with apparent negative effects on the plant reproduction; and (iv) the close relationship between H. aristata and carpenter bees was in agreement with the large-bee pollination syndrome of this plant. Our results highlight the need for studies detecting the roles of individual visitors. We showed that such an approach is necessary to evaluate the pollination syndrome hypothesis and create relevant evolutionary and ecological hypotheses.     

Negative native–exotic diversity relationship in oak savannas explained by human influence and climate

Recent research has proposed a scale-dependence to relationships between native diversity and exotic invasions. At fine spatial scales, native–exotic richness relationships should be negative as higher native richness confers resistance to invasion. At broad scales, relationships should be positive if natives and exotics respond similarly to extrinsic factors. Yet few studies have examined both native and exotic richness patterns across gradients of human influence, where impacts could affect native and exotic species differently. We examined native–exotic richness relationships and extrinsic drivers of plant species richness and distributions across an urban development gradient in remnant oak savanna patches. In sharp contrast to most reported results, we found a negative relationship at the regional scale, and no relationship at the local scale. The negative regional-scale relationship was best explained by extrinsic factors, surrounding road density and climate, affecting natives and exotics in opposite ways, rather than a direct effect of native on exotic richness, or vice versa. Models of individual species distributions also support the result that road density and climate have largely opposite effects on native and exotic species, although simple life history traits (life form, dispersal mode) do not predict which habitat characteristics are important for particular species. Roads likely influence distributions and species richness by increasing both exotic propagule pressure and disturbance to native species. Climate may partially explain the negative relationship due to differing climatic preferences within the native and exotic species pools. As gradients of human influence are increasingly common, negative broad-scale native–exotic richness relationships may be frequent in such landscapes.     

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.     

Pollination Ecology and Seed Production of Rhododendron Ponticum in Native and Exotic Habitats

Alien plants may be reproductively limited in exotic habitats because of a lack of mutualistic pollinators. However, if plants are adequately served by generalist pollinators, successful reproduction, naturalisation and expansion into exotic habitats may occur. Rhododendron ponticum is very successful, ecologically damaging invasive plant in Britain and Ireland, but is in decline in its native Iberian habitat. It spreads locally by sending out lateral branches, but for longer distance dispersal it relies on sexually produced seeds. Little is known about R. ponticum's pollination ecology and breeding biology in invaded habitats. We examined the flower-visiting communities and maternal reproductive success of R. ponticum in native populations in southern Spain and in exotic ones in Ireland. R. ponticum in flowers are visited by various generalist (polylectic) pollinator species in both native and exotic habitats. Although different species visited flowers in Ireland and Spain, the flower visitation rate was not significantly different. Insects foraging on R. ponticum in Spain carried less R. ponticum pollen than their Irish counterparts, and carried fewer pollen types. Fruit production per inflorescence varied greatly within all populations but was significantly correlated with visitation at the population level. Nectar was significantly depleted by insects in some exotic populations, suggesting that this invasive species is providing a floral resource for native insects in some parts of Ireland. The generality of the pollination system may be factor contributing to R.ponticum's success in exotic habitats.     

Comparisons of arthropod assemblages on an invasive and native trees: abundance,diversity and damage

The success of exotic plants may be due to lower herbivore loads than those on native plants (Enemies Release Hypothesis). Predictions of this hypothesis include lower herbivore abundances, diversity, and damage on introduced plant species compared to native ones. Greater density or diversity of predators and parasitoids on exotic versus native plants may also reduce regulation of exotic plants by herbivores. To test these predictions, we measured arthropod abundance, arthropod diversity, and foliar damage on invasive Chinese tallow tree (Triadica sebifera) and three native tree species: silver maple (Acer saccharinum), sycamore (Platanus occidentalis), and sweetgum (Liquidambar styraciflua). Arthropod samples were collected with canopy sweep nets from six 20 year old monoculture plots of each species at a southeast Texas site. A total of 2,700 individuals and 285 species of arthropods were caught. Overall, the species richness and abundance of arthropods on tallow tree were similar to the natives. But, ordination (NMS) showed community composition differed on tallow tree compared to all three native trees. It supported an arthropod community that had relatively lower herbivore abundance but relatively more predator species compared to the native species examined. Leaves were collected to determine damage. Tallow tree experienced less mining damage than native trees. The results of this study supported the Enemies Release Hypothesis predictions that tallow tree would have low herbivore loads which may contribute to its invasive success. Moreover, a shift in the arthropod community to fewer herbivores without a reduction in predators may further limit regulation of this exotic species by herbivores in its introduced range.     

Comparison of native and exotic distribution and richness models across scales reveals essential conservation lessons

Comparing native and exotic plant species distribution and richness models can help to reveal the causes of invasive exotic species proliferation and provide recommendations for preserving native‐dominated ecosystems. However, models may have limited applicability if potentially divergent patterns across scales, spatial autocorrelation and correspondence with community‐wide patterns such as species richness are not considered. I modeled the distributions of 20 dominant native and 20 dominant exotic species among and within patches in a heavily‐invaded and threatened ecosystem in western North America, examining the roles of scale and species origin on variable selection, spatial autocorrelation and model accuracy to determine conditions that favour native over exotic dominants, and derive recommendations for effective management. I also compared distribution models with native and exotic species richness models, to determine the extent to which dominant native and exotic species were representative of synoptic community patterns. Predictability was lower for exotic dominants, possibly because they are environmental generalists, and was lower within than among patches. Predictors were generally shared between distribution and richness models; however, species‐specific differences were common within both native and exotic species groups. Predictors for individual species across scales were frequently different and sometimes opposing. Distribution and richness models suggest that management assuming environmental affiliation at one scale may be ineffective at another; that site prioritization to maximize native versus exotic richness may not preserve the habitat of some common native species; and that intensive management to reduce exotics may be difficult due to low predictability and shared affiliations with natives. Comparing native and exotic distribution and richness models at two scales enabled scale‐specific conservation recommendations and elucidated trade‐offs between management for richness and representation that distribution models at an individual scale would not have allowed.     

Native versus exotic community patterns across three scales: Roles of competition,environment and incomplete invasion

Three fundamental, interrelated questions in invasion ecology are: (1) to what extent do exotic species outcompete natives; (2) are native and exotic communities functionally similar or different; and (3) are differences in biogeographic patterns in native and exotic communities due to incomplete invasions among exotics? These questions are analogous to general questions in community ecology regarding the relative roles of competition, environmental response and dispersal limitation in community assembly. We addressed each of these questions for plant communities in discrete meadow patches, using analyses at three scales ranging from the landscape to microsites. A weak positive relationship between native and exotic species richness in microsites, and a predominance of positive correlations in abundance among native and exotic species pairs suggest that competition has been less important than other factors in determining native versus exotic abundance and community composition. In contrast, models of species richness and community compositional change across scales suggest native versus exotic community patterns are largely determined by a mix of scale-dependent concordant (shared positive or negative) and discordant relationships with environmental variables. In addition, detailed analyses of species-area and species-abundance relationships suggest ongoing expansion of exotic species populations, indicating that the assembly of the exotic community is in its early stages. Thus, while competition does not appear to strongly affect native versus exotic abundances and compositions at present, it may intensify in the future. Our results indicate that synoptic patterns in native versus exotic richness that have been previously attributed to a single cause may in fact be due to a complex mix of concordant and discordant responses to environmental factors across scales. They also suggest that conservation efforts aimed at promoting natives and reducing exotics should focus on the factors and scales for which such a response (i.e., promotion of high native and low exotic richness) can be expected.