Causes of exotic bird establishment across oceanic islands

The probability that exotic species will successfully establish viable populations varies between regions, for reasons that are currently unknown. Here, we use data for exotic bird introductions to 41 oceanic islands and archipelagos around the globe to test five hypotheses for this variation: the effects of introduction effort, competition, predation, human disturbance and habitat diversity (island biogeography). Our analyses demonstrate the primary importance of introduction effort for avian establishment success across regions, in concordance with previous analyses within regions. However, they also reveal a strong negative interaction across regions between establishment success and predation; exotic birds are more likely to fail on islands with species-rich mammalian predator assemblages.     

Biogeography of grasses (Poaceae) on islands of southwestern Australia

Abstract The number of native grass species and exotic grass species present on 129 offshore islands of southwestern Australia is best predicted by island area and island disturbance, respectively. Isolation of islands and gull activity on islands only slightly improved these predictions. Species turnover on a subset of 30 islands indicated that exotic grass species were more prone to local extinction and more likely to immigrate than native grass species. The major conservation implication of this study is that habitat disturbance on these islands should be minimized to reduce establishment of exotic grass species.     

Does Invasion of Exotic Plants Promote Invasion of Exotic Flower Visitors? A Case Study from the Temperate Forests of the Southern Andes

Habitat disturbance, particularly of human origin, promotes the invasion of exotic plants, which in turn might foster the invasion of alien-interacting animals. Here we assess whether the invasion of exotic plants – mostly mediated by habitat disturbance – facilitates the invasion of exotic flower visitors in temperate forests of the southern Andes, Argentina. We recorded visit frequencies and the identity of visitors to the flowers of 15 native and 15 exotic plant species occurring in different highly disturbed and less disturbed habitats. We identified three alien flower visitors, the hymenopterans Apis mellifera, Bombus ruderatus, and Vespula germanica. We found significantly more visitation by exotic insects in disturbed habitats. This pattern was explained, at least in part, by the association between alien flower visitors and flowers of exotic plants, which occurred more frequently in disturbed habitats. However, this general pattern masked different responses between the two main alien flower visitors. Apis mellifera exploited almost exclusively the flowers of a subset of herbaceous exotic plants that thrive under disturbance, whereas B. ruderatus visited equally flowers of both exotic and native plants in both disturbed and undisturbed habitats. We did not find any strong evidence that flowers of exotic plants were more generalist than those of native plants, or that exotic flower visitors were more generalist than their native counterparts. Our results suggest that alien plant species could facilitate the invasion of at least some exotic flower visitors to disturbed habitats. Because flowering plants as well as flower visitors benefit from this mutualism, this association may enhance, through a positive feedback, successful establishment of both exotic partners.     

Biological soil crusts determine the germination and growth of two exotic plants

In arid and semiarid ecosystems, the potential threats of exotic invasive species are enhanced due to increasing human activities. Biological soil crusts (BSCs), acting as arid ecosystem engineers, may play an important role in preventing the establishment of exotic invasive plants. Our goal was to examine whether BSCs could inhibit the establishment of probable exotic plant species originating from adjacent grasslands located along the southeast edge of the Tengger Desert. In our study, we investigated the effects of three BSC types (cyanobacteria, lichen, and moss crusts) under two disturbance conditions (intact and disturbed) on the establishment of two exotic plant species (Ceratoides latens and Setaria viridis) using indoor experiments. We found both negative and positive effects of BSCs on the establishment of the two exotic plant species. Compared with the disturbed BSCs, the germination percentages of C. latens and S. viridis were reduced by 54% to 87% and 89% to 93%, respectively, in intact BSCs. In contrast, BSCs significantly promoted the height growth and aboveground biomass of the two exotic plant species (p < .05) by enhancing the soil water and nutrient availability for the exotic plants. Our results confirm that BSCs strongly suppress the rapid expansion of exotic plant populations by inhibiting germination of seed with big size or appendages and have a weak inhibitory effect on exotic plant with small and smooth seeds. This may decrease the threat of propagation of exotic species. In the meantime, BSCs promote the growth of a few successful engraftment seedlings, which increased the beta diversity. Our work suggests that better understanding the two opposing effects of BSCs on the establishment of exotic plant species in different growth stages (germination and growth) is important for maintaining the health and stability of revegetated regions.     

Exotic vascular plant invasiveness and forest invasibility in urban boreal forest types

The riverine forests of the northern city of Edmonton, Alberta, Canada display strong resilience to disturbance and are similar in species composition to southern boreal mixedwood forest types. This study addressed questions such as, how easily do exotic species become established in urban boreal forests (species invasiveness) and do urban boreal forest structural characteristics such as, native species richness, abundance, and vertical vegetation layers, confer resistance to exotic species establishment and spread (community invasibility)? Eighty-four forest stands were sampled and species composition and mean percent cover analyzed using ordination methods. Results showed that exotic tree/shrub types were of the most concern for invasion to urban boreal forests and that exotic species type, native habitat and propagule supply may be good indicators of invasive potential. Native forest structure appeared to confer a level of resistance to exotic species and medium to high disturbance intensity was associated with exotic species growth and spread without a corresponding loss in native species richness. Results provided large-scale evidence that diverse communities are less vulnerable to exotic species invasion, and that intermediate disturbance intensity supports species coexistence. From a management perspective, the retention of native species and native forest structure in urban forests is favored to minimize the impact of exotic species introductions, protect natural succession patterns, and minimize the spread of exotic species.     

The diversity and origin of exotic ants arriving in New Zealand via human-mediated dispersal

The number of exotic ant species being dispersed to new regions by human transportation and the trade pathways responsible for this are poorly understood. In this study, the taxonomic diversity, trade pathways, and origin of exotic ants intercepted at the New Zealand border were examined for the period 1955–2005. Overall, there were a total 4355 interception records, with 115 species from 52 genera. The 10 most frequently intercepted genera, and the 20 most frequently intercepted species contributed > 90% of all records. Many of the species frequently intercepted are regarded as invasive species, and several are established in New Zealand. The most intercepted species was Pheidole megacephala . Despite a relatively low trade relationship, a high proportion (> 64%) of the exotic ants which were intercepted originated from the Pacific region. However, the majority of species intercepted from the Pacific was exotic to the region (71%), or to a lesser extent, wide-ranging Pacific native species. No endemic species from the Pacific were intercepted. The effectiveness of detecting exotic ant species at the New Zealand border ranged from 48–78% for different trade pathways, indicating a number of species remain undetected. Trade routes associated with specific geographical regions represent a major filter for the arrival of exotic ant species. Despite the importance of the Pacific as a frequent pathway, we suggest that the future establishment of exotic ant species in New Zealand is likely to be mitigated by a renewed focus on trade routes with cool temperate regions, particularly Australia.     

Takahe Valley Hut: a focal point for weed invasion in an isolated area of Fiordland National Park, New Zealand

The role of backcountry huts as focal points for weed establishment and spread into New Zealand’s national parks has received little attention. In this study we describe the pattern of weed spread around Takahe Valley Hut, Murchison Mountains, Fiordland National Park. Established in 1948, the hut is located at 900 m a.s.l. at the ecotone between Nothofagus forest and valley floor shrubland/grassland. We recorded the distribution of vascular plants in quadrats (110) placed by restricted randomisation around the hut, and measured relative irradiance and distance from the hut. Nine exotic species, mostly grasses, were recorded, the most frequent being Agrostis capillaris (34%). The majority of occurrences of exotic plant species were located in the immediate vicinity (less than 5 m) of the hut but two exotic species (Agrostis capillaris and Dactylis glomerata) ranged more widely. Exotic species were present in well-drained shrubland and grassland but did not extend far into Nothofagus forest or onto infertile wetlands. The percentage of exotic species in quadrats declined significantly with distance from the hut. There was no linear relationship between the percentage of exotic species and relative irradiance. When forest quadrats were excluded, the number of native species in quadrats was negatively correlated with the number of exotic species, suggesting competitive displacement of native species by exotics in non-forest habitats. Long-term persistence of most exotic species at this site depends on physical disturbance and nutrient enrichment associated with human activities at the hut site. However the maintenance of this species pool has provided sufficient propagule pressure for some exotic species to disperse into the wider area. Weed accumulation around huts can be reduced by locating huts in vegetation types that are more resistant to invasion, and maintaining facilities to eliminate local weed infestations.     

Establishment of exotic parasites: the origins and characteristics of an avian malaria community in an isolated island avifauna

Knowledge of the processes favouring the establishment of exotic parasites is poor. Herein, we test the characteristics of successful exotic parasites that have co-established in the remote island archipelago of New Zealand, due to the introduction of numerous avian host species. Our results show that avian malaria parasites (AM; parasites of the genus Plasmodium) that successfully invaded are more globally generalist (both geographically widespread and with a broad taxonomic range of hosts) than AM parasites not co-introduced to New Zealand. Furthermore, the successful AM parasites are presently more prevalent in their native range than AM parasites found in the same native range but not co-introduced to New Zealand. This has resulted in an increased number and greater taxonomic diversity of AM parasites now in New Zealand.     

Invasive autumn olive performance varies in different reclamation conditions: implications for restoration

Surface mining has caused significant disturbance globally, and is responsible for the loss of more than 600,000 ha of the world's largest temperate deciduous forest in the Appalachian region of the United States alone. Due to the heavy disturbance on mine lands, invasive plants have become dominant on many former coalfields, some of which were intentionally planted with exotic species. The establishment of invasive plants on these disturbed lands has often led to reductions in establishment of desirable native species. Autumn olive (Elaeagnus umbellata), an exotic nitrogen‐fixing shrub, is a problematic invasive species on reclaimed sites in Appalachia. To better understand how reclamation conditions affect autumn olive, we assessed the effects of the mine‐soil substrate and vegetation seeding on autumn olive establishment and growth. In each experiment, we also manipulated the herbaceous plant community to further examine effects on autumn olive establishment and growth. In spring 2015, we transplanted 480 1‐year‐old autumn olive seedlings across both experiments. After 2 years of growth, autumn olive performed better in weathered‐rock than in unweathered‐rock substrates; in bare‐ground plots than in vegetated plots; and in tree‐compatible (low‐competition) seeding than in more‐competitive conventional seeding. No treatment precluded autumn olive establishment. However, our results show that strategic use of beginning substrates and planting mixes can have strong inhibitory effects on invasive plants, but also that substrate and herbaceous‐plant community conditions favorable to establishment of native trees are also favorable to autumn olive.     

Management strategies for plant invasions: manipulating productivity, disturbance, and competition

The traditional approach to understanding invasions has focused on properties of the invasive species and of the communities that are invaded. A well‐established concept is that communities with higher species diversity should be more resistant to invaders. However, most recently published field data contradict this theory, finding instead that areas with high native plant diversity also have high exotic plant diversity. An alternative environment‐based approach to understanding patterns of invasions assumes that native and exotic species respond similarly to environmental conditions, and thus predicts that they should have similar patterns of abundance and diversity. Establishment and growth of native and exotic species are predicted to vary in response to the interaction of plant growth rates with the frequency and intensity of mortality‐causing disturbances. This theory distinguishes between the probability of establishment and the probability of dominance, predicting that establishment should be highest under unproductive and undisturbed conditions and also disturbed productive conditions. However, the probability of dominance by exotic species, and thus of potential negative impacts on diversity, is highest under productive conditions. The theory predicts that a change in disturbance regime can have opposite effects in environments with contrasting levels of productivity. Manipulation of productivity and disturbance provides opportunities for resource managers to influence the interactions among species, offering the potential to reduce or eliminate some types of invasive species.     

SNP analyses reveal a diverse pool of potential colonists to earthquake‐uplifted coastlines

In species that form dense populations, major disturbance events are expected to increase the chance of establishment for immigrant lineages. Real‐time tests of the impact of disturbance on patterns of genetic structure are, however, scarce. Central to testing these concepts is determining the pool of potential immigrants dispersing into a disturbed area. In 2016, a 7.8 magnitude earthquake occurred on the South Island of New Zealand. Affecting approximately 100 km of coastline, this quake caused extensive uplift (several metres high), extirpating many intertidal populations, including keystone intertidal kelp species. Following the uplift, we set out to determine the geographic origins of detached kelp specimens which rafted into the disturbed zone. Specifically, we used genotyping‐by‐sequencing (GBS) approaches to compare beach‐cast southern bull‐kelp (Durvillaea antarctica and Durvillaea poha) samples to established populations throughout the species' ranges, and thus infer the geographic origins of potential colonists reaching the disturbed coast. Our findings revealed an ongoing supply of diverse lineages dispersing to the newly uplifted coastline, suggesting potential for establishment of “exotic” lineages following disturbance. Furthermore, we found that some drifting individuals of each species came from far‐distant regions, some >1,200 km away. These results show that diverse lineages – in many cases from very distant sources – can compete for new space in the wake of an exceptional disturbance event, illustrating the potential of long‐distance dispersal as a key mechanism for reassembly of coastal ecosystems. Furthermore, our findings demonstrate that high‐resolution genomic baselines can be used to robustly assign the provenance of dispersing individuals.     

Susceptibility of A Patagonian Mallín Flooded Meadow to Invasion by Exotic Species

Mallín wetland meadows are highly diverse, rare habitats in western Patagonia that are believed to be particularly susceptible to disturbance. I tested the hypothesis that exotic species reduce the rate and extent of recolonization by native species after disturbances. Open patches (50 × 50 cm) were subjected to artificial seeding with propagules of two exotic species. Cover of individual species in each plot was measured over the four following austral summers (1990–1993) and two late summers (1995 and 1998). The effect of a particular exotic species was not the same for all native species. For some native species, the presence of an exotic species in the plots was associated with a significant increase in cover, while a significant decrease in cover was observed for other species. Native species richness and diversity were not significantly affected by the introduction and establishment of exotic species. In addition, the failure of the exotic species to establish in the undisturbed control plots further suggests that the undisturbed mallín is resistant to invasion by exotic species. The results suggest that the small-scale changes that occurred in this community due to exotic species during recolonization were transient and almost undetectable after eight years.     

Covariance between disturbance and soil resources dictates the invasibility of northern fescue prairies

Increasing environmental impacts of exotic organisms have refocused attention on the ability of diverse communities to resist biological invaders. Although resource availability, often related to natural and anthropogenic disturbances, appears central to the invasibility of biological communities, understanding the links between resources, diversity and invasibility is often confounded by the covariance among key variables. To test the hypothesis that community invasibility remains contingent on the type and intensity of disturbance and their impacts on plant community diversity and resource availability, we designed an experiment testing the invasibility of northern fescue prairies by smooth brome (Bromus inermis Leyss.), a Eurasian perennial grass, threatening the structure and function of prairie remnants throughout the Great Plains. Using soil disturbances and herbicide, we imposed treatments manipulating the diversity and resource availability of native prairies. Our observations demonstrate that the vulnerability of native prairies to exotic plant invasions remains contingent on resources. While the establishment of smooth brome seedlings increased with increasing disturbance, its impact depended on the availability of soil nitrogen. As a result, soil burial treatments, simulating disturbance by northern pocket gophers, provided poor recruitment areas for smooth brome, and their low levels of soil moisture and nitrogen, combined with the rapid recovery of the prairie community, compromised seedling establishment. Emphasizing the covariance of diversity and key environmental variables following disturbance, our findings illustrate the importance of disturbance type and intensity on community invasibility. Such a consideration is critical in the conservation and restoration of native prairie remnants throughout the Great Plains.     

Biological Invasions in the Amazonian Tropical Rain Forest: The Case of Drosophilidae (Insecta,Diptera) in Ecuador,South America

There have been few reports of invasions in continental rain forests, especially for exotic animals. This study provides original data concerning the potential of exotic drosophilid species to colonize the Amazonian tropical rain forest. To investigate if the structure of drosophilid assemblages differed in response to anthropogenic disturbance, we performed a taxonomic survey at six sites within the Yasuni National Park in Ecuadorian Amazonia along a disturbance gradient from pristine to clearcut artificial forest. A total of 7425 individuals from 34 species were collected of which seven species were exotic. There was significant variation in the assemblage composition along this disturbance gradient; 31 percent of which was explained by the presence of exotic species, particularly at the most disturbed sites, through nonmetric multidimensional scaling and SIMPER analyses. These results confirm the susceptibility of continental rain forests to invasion by exotic species. There is an urgent need to develop and implement monitoring systems, for example, based on drosophilid assemblage surveys, to detect exotic invasions in continental tropical forests.     

Does disturbance,competition or resource limitation underlie Hieracium lepidulum invasion in New Zealand? Mechanisms of establishment and persistence,and functional differentiation among invasive and native species

The processes underlying plant invasions have been the subject of much ecological research. Understanding mechanisms of plant invasions are difficult to elucidate from observations, yet are crucial for ecological management of invasions. Hieracium lepidulum, an asteraceous invader in New Zealand, is a species for which several explanatory mechanisms can be raised. Alternative mechanisms, including competitive dominance, disturbance of resident vegetation allowing competitive release or nutrient resource limitation reducing competition with the invader are raised to explain invasion. We tested these hypotheses in two field experiments which manipulated competitive, disturbance and nutrient environments in pre‐invasion and post‐invasion vegetation. H. lepidulum and resident responses to environmental treatments were measured to allow interpretation of underlying mechanisms of establishment and persistence. We found that H. lepidulum differed in functional response profile from native species. We also found that other exotic invaders at the sites were functionally different to H. lepidulum in their responses. These data support the hypothesis that different invaders use different invasion mechanisms from one another. These data also suggest that functional differentiation between invaders and native resident vegetation may be an important contributing factor allowing invasion. H. lepidulum appeared to have little direct competitive effect on post‐invasion vegetation, suggesting that competition was not a dominant mechanism maintaining its persistence. There was weak support for disturbance allowing initial establishment of H. lepidulum in pre‐invasion vegetation, but disturbance did not lead to invader dominance. Strong support for nutrient limitation of resident species was provided by the rapid competitive responses with added nutrients despite presence of H. lepidulum. Rapid competitive suppression of H. lepidulum once nutrient limitation was alleviated suggests that nutrient limitation may be an important process allowing the invader to dominate. Possible roles of historical site degradation and/or invader‐induced soil chemical/microbial changes in nutrient availability are discussed.     

Challenges and limitations to native species restoration in the Great Basin,USA

The Great Basin of the western USA is an arid region characterized by high spatial and temporal variability. The region experienced high levels of ecological disturbance during the early period of Euro-American settlement, especially from about 1870–1935. The principal plant communities of the Great Basin are sagebrush steppes, dominated by various Artemisia shrubs and perennial bunchgrasses that represent the largest rangeland ecosystem in North America. In low to mid-elevation sagebrush communities, exotic annual grasses have displaced native plant species and are associated with a dramatic increase in size and frequency of wildfires. Degradation in this region is driven by processes that cause the loss of mature bunchgrasses, which, when intact, limit exotic annual grass invasion. Historically, large economic investments to restore degraded Great Basin rangeland through establishment of native bunchgrasses, principally utilizing heavily mechanized agronomic approaches, have been met with limited success. A multitude of environmental factors contribute to the lack of restoration success in this region, but seedling mortality from freezing and drought has been identified as a primary demographic limitation to successful bunchgrass establishment. Novel approaches to overcoming limitations to bunchgrass establishment will be required for restoration success. Increased national concern and a near listing of the greater sage-grouse, a steppe-obligate species, to Endangered Species status, has spurred greater regional support and collaboration across a diversity of stakeholder groups such as state and federal land and wildlife management agencies, county planners, and ranchers.     

Effects of grazing on seedling establishment: the role of seed and safe-site availability

Abstract. The first objective of this paper was to assess the effects of grazing on seedling establishment of two species whose relative abundance at the adult stage is affected by grazing in a contrasting fashion. Second, we evaluated the relative importance of seed versus safe-site availability in explaining the effect of grazing on seedling establishment. We monitored seedling establishment on a grazed area, on two areas which had not been grazed for two and seven years, and on plots which had been experimentally defoliated. The species compared were Dan-thonia montevidensis, a native perennial grass which dominates both grazed and ungrazed communities, and Leontodón taraxacoides, an invading exotic rosette species from the Compositae family. Continuous grazing enhanced seedling establishment of both species through its effect on the availability of safe sites. Seed availability accounted for only one, but very important, grazing effect: the lack of response by L. taraxacoides to the defoliation in the seven-year old exclosure. Its seed supply was depleted by exclusion of grazing and, consequently, its short-term regeneration capacity after disturbance was lost.     

Relationships between anthropogenic disturbance,soil properties and plant invasion in endangered Cumberland Plain Woodland,Australia

Abstract Invasive exotic plants are a significant threat to areas of conservation value, with endangered ecological communities being especially vulnerable. We assessed the role of different anthropogenic disturbances in determining the success of exotic plants in the endangered Cumberland Plain Woodland community of western Sydney and examined the impact of these disturbances on soil characteristics that are likely to impact on vegetation, including total P, pH, water retention capacity, organic matter content and electrical conductivity. The disturbance types were: (i) land use incorporating clearing, agriculture and grazing by stock; (ii) creeks draining a developed catchment; and (iii) roads. Remnants that had been cleared and grazed had higher exotic and lower native species richness and cover than all other disturbance types. Areas that had been grazed but not cleared did not have more exotic species richness or cover than uncleared/ungrazed areas, thus retaining high conservation value. Areas within 2 m of a creek edge had higher exotic species richness and cover than areas further from the creek edge. Areas downslope of sealed roads had significantly higher exotic species richness and cover than areas below unsealed roads. No single soil attribute or combination of soil attributes was consistently able to account for variation in exotic species cover under the different disturbance types. Thus it appears that other factors such as site history and propagule pressure may be more important in determining exotic species success than soil characteristics alone, in this vegetation community.     

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.     

Rare species loss alters ecosystem function – invasion resistance

The imminent decline in species diversity coupled with increasing exotic species introductions has provoked investigation into the role of resident diversity in community resistance to exotic species colonization. Here we present the results of a field study using an experimental method in which diversity was altered by removal of less abundant species and the resulting disturbance was controlled for by removal of an equivalent amount of biomass of the most common species from paired plots. Following these manipulations, the exotic grass, Lolium temulentum , was introduced. We found that exotic species establishment was higher in plots in which diversity was successfully reduced by removal treatments and was inversely related to imposed species richness. These results demonstrate that less common species can significantly influence invasion events and highlight the potential role of less common species in the maintenance of ecosystem function.