Grass invasion causes rapid increases in ecosystem carbon and nitrogen storage in a semiarid shrubland

Accurately predicting terrestrial carbon (C) and nitrogen (N) storage requires understanding how plant invasions alter cycling and storage. A common, highly successful type of plant invasion occurs when the invasive species is of a distinctly different functional type than the native dominant plant, such as shrub encroachment throughout the western United States and annual grass invasions in Mediterranean shrublands, as studied here. Such invasions can dramatically transform landscapes and have large potential to alter C and N cycling by influencing storage in multiple pools. We used a manipulation of non‐native annual grass litter within a shrub‐dominated habitat in southern California (coastal sage scrub, CSS) to study how grass invasion alters ecosystem C and N storage. We added, removed, or left unchanged grass litter in areas of high and low invasion, then followed soil and vegetation changes. Grass litter greatly increased C and N storage in soil, aboveground native and non‐native biomass. Aboveground litter storage increased due to the greater inputs and slower decomposition of grass litter relative to shrub litter; shading by grass litter further reduced decomposition of both non‐native and native litter, which may be due to reduced photodegradation. Soil C and N pools in areas of high litter increased ～20% relative to low litter areas in the two years following manipulation and were generally sinks for C and N, while areas with low litter were sources. We synthesize our results into a C cycle of invaded and uninvaded areas of CSS and link changes in storage to increases in the soil fungi : bacteria ratio, increased plant inputs, and decreased litter loss. Overall, we show that grasses, especially through their litter, control important abiotic and biotic mechanisms governing C and N storage, with widespread implications for C sequestration and N storage in semiarid systems undergoing grass or shrub invasions.     

Invasive grass litter facilitates native shrubs through abiotic effects

Questions: Plant invasions are considered one of the top threats to the biodiversity of native taxa, but clearly documenting the causal links between invasions and the decline of native species remains a major challenge of invasion biology. Most studies have focused on impacts of invaders' living biomass, rather than on mechanisms mediated by litter. However, invasive plant litter, which is often of a very different type and quantity than a system's native plant litter, can have multiple important effects on ecosystem processes – such as nitrogen cycling and soil microclimate – that may influence native plants. Location: We studied effects of litter of invasive grass species that are widespread throughout western North America on native shrubs in southern California's semi‐arid habitat of coastal sage scrub. Methods: We combined a 3‐year field manipulation of non‐native litter with structural equation modeling to understand interacting effects on non‐native grasses, native shrubs, soil nitrogen (available and total), and soil moisture. Results: Litter addition facilitated non‐native grass growth, revealing a positive feedback likely to enhance invasion success. Contrary to a major paradigm of invasion biology – that competition with invasive plant species causes declines of native plants – we found that litter also facilitated growth of the native dominant shrub, a result supported by observational trends. Structural equation models indicated that enhanced soil moisture mediated the positive effects of litter on shrub growth. Conclusions: We demonstrate that invasive plants, via their litter, can facilitate dominant native plants by altering soil moisture. Our results highlight that understanding the impacts and mechanisms of plant invasions may be enhanced by considering the role of invasive plant litter on native plants and ecosystem properties.     

The effects of development,vegetation‐type conversion,and fire on low‐elevation Southern California spider assemblages

California sage scrub (CSS), a native ecosystem type of low‐elevation areas of Southern California, is increasingly threatened by urban development, altered fire regimes, and vegetation‐type conversion to non‐native grasslands. Using pitfall traps, we examined how suburbanization, type conversion, and fire influence ground‐dwelling spider assemblages in eastern Los Angeles County, CA, by surveying spiders in three habitats (CSS, non‐native grasslands, and suburban areas) before and after a fire that occurred in a small portion of our study site. Spider assemblages in the suburban habitat differed from those in CSS and non‐native grassland habitats, but CSS and grassland assemblages did not significantly differ. This suggests that the urban development, but not vegetation‐type conversion to non‐native grasslands, has significant effects on ground‐dwelling spider assemblages. Fire had no observable effect on assemblages. Because ground‐dwelling spiders were not impacted by fire and type conversion, increased fire frequencies, which often result in the establishment of non‐native grasses, may not deleteriously influence this animal group, a differing pattern from other taxonomic groups. However, the rapid urban development occurring in low‐elevation areas of Southern California means that species requiring non‐suburban sites for their survival (15 species, 24.1%) may be threatened and require conservation assessment.     

Granivory in California sage scrub: implications for common plant invaders and ecosystem conservation

Seed predation may influence community assembly and invasion dynamics when seed predators preferentially select some seed species over others. However, the role of different seed predators and their preferences for seeds in multiple ecological contexts, including the endangered California sage scrub ecosystem, remain unresolved making predictions about which processes limit or promote invasions difficult. In addition, selection criteria, specifically how seed size and biogeographic origin (native versus invasive) influences selection, requires further research across multiple granivore guilds. We examined seed predator preference for common native and invasive seeds across small and large seed classes using seed dish experiments with motion-sensor video observation. We also quantified the relative importance of ants, birds, and mammals as seed predators in sage scrub. Community-wide, granivores preferred the small-seeded invasive mustard Brassica nigra, avoided the large-seeded invasive grass Bromus rubens and the native large-seeded shrub Encelia californica, and did not show significant selection or avoidance for the small-seeded native shrub Salvia apiana. Birds, notably California towhees (Melozone crissalis), were the most frequent granivore while rodents and ants rarely removed seeds. This study reveals that birds are important seed predators in California sage scrub and have the potential to contribute to biotic resistance to mustard invasions (Brassica nigra and potentially others).

     

Restoration of a Native Shrubland Impacted by Exotic Grasses, Frequent Fire, and Nitrogen Deposition in Southern California

Natural ecosystems globally are often subject to multiple human disturbances that are difficult to restore. A restoration experiment was done in an urban fragment of native coastal sage scrub vegetation in Riverside, California that has been subject to frequent fire, high anthropogenic nitrogen deposition, and invasion by Mediterranean annual weeds. Hand cultivation and grass‐specific herbicide were both successful in controlling exotic annual grasses and promoting establishment of seeded coastal sage scrub vegetation. There was no native seedbank left at this site after some 30 years of conversion to annual grassland, and the only native plants that germinated were the seeded shrubs, with the exception of one native summer annual. The city green‐waste mulch used in this study (C:N of 39:1) caused short‐term N immobilization but did not result in decreased grass density or increased native shrub establishment. Seeding native shrubs was successful in a wet year in this Mediterranean‐type climate but was unsuccessful in a dry year. An accidental spring fire did not burn first‐year shrubs, although adjacent plots dominated by annual grass did burn. The shrubs continued to exclude exotic grasses into the second growing season, suggesting that successful shrub establishment may reduce the frequency of the fire return interval.     

Aphid‐tending ants on introduced fennel: can resources derived from non‐native plants alter the trophic position of higher‐order consumers?

1. Although plant invasions often reduce insect abundance and diversity, non‐native plants that support phytophagous insects can subsidise higher trophic levels via elevated herbivore abundance. 2. Here ant–aphid interactions on non‐native fennel on Santa Cruz Island, California are examined. Fennel hosts abundant, honeydew‐producing fennel aphids. The patchiness of fennel and the relative lack of honeydew‐producing insects on other plants at our study sites suggest that assimilation of fennel‐derived honeydew would increase the abundance and decrease the trophic position of the omnivorous, aphid‐tending Argentine ant. 3. To assess the strength of the ant–aphid interaction, a comparison of ant abundance on and adjacent to fennel prior to and 3 weeks after experimental aphid removal was performed. Compared with control plants with aphids, ants declined in abundance on and around fennel plants following aphid removal. At the habitat scale, pitfall traps in fennel‐dominated habitats captured more ants than in fennel‐free scrub habitats. 4. To determine if assimilation of aphid‐produced honeydew reduces the ant's trophic position, variation in δ¹⁵N values among ants, plants and other arthropods was analysed. Unexpectedly, δ¹⁵N values for ants in fennel‐dominated habitats were higher than those of arthropod predators from the same sites and also higher than those of ants from fennel‐free habitats. 5. Our results illustrate how introduced plants that support phytophagous insects appear to transfer energy to higher trophic levels via elevated herbivore abundance. Although assimilation of fennel‐derived honeydew did not appear to reduce consumer trophic position, spatial variation in alternative food resources might obscure contributions from honeydew.     

Does the proportion of arthropods versus fruit in the diet influence overwintering condition of an omnivorous songbird?

Many migratory songbirds switch from a primarily insectivorous diet during the breeding season to either a mixed diet or fruit diet during the non‐breeding season. However, for species with mixed diets, arthropods may be superior food items because of their higher protein content and easier digestibility. We tested this hypothesis by analyzing the diet and body condition of omnivorous Wood Thrushes (Hylocichla mustelina) at a non‐breeding site in tropical forest in Belize, Central America. We used analysis of stable isotopes δ¹⁵N and δ¹³C in the blood to measure diet. Our objective was to determine if a higher dietary proportion of arthropods relative to fruit (i.e., higher δ¹⁵N and δ¹³C) was associated with better body condition. We also examined the possible effect of age, sex, and habitat type on Wood Thrush diets, as well as any changes in diet through the overwintering period. We used a hierarchical Bayesian mixing model (MixSIAR) to estimate the proportion of different prey items in the diet of overwintering Wood Thrushes overall, in each habitat type, and over time during the non‐breeding period. From January to April, we found a significant decline in δ¹⁵N in forest habitats, whereas δ¹⁵N increased in scrub habitat. There was no significant seasonal change in δ¹³C. Birds with higher δ¹⁵N or δ¹³C values were not in better body condition. Females in dry‐scrub habitat consumed more fruit than males, but this did not affect body condition. Mixing model results indicated that most Wood Thrushes at our study sites consumed primarily arthropods, even during the driest times of the non‐breeding season and in the driest habitat. Overall, our results suggest that the diet of Wood Thrushes varies with habitat and during the overwintering period, but diet alone was not a predictor of body condition. Wood Thrushes, and possibly other omnivorous migratory songbirds, are apparently flexibly able to meet their wintering and pre‐migration nutritional demands with a variety of diets.     

The distribution and habitat associations of non‐native plant species in urban riparian habitats

Questions: 1. What are the distribution and habitat associations of non‐native (neophyte) species in riparian zones? 2. Are there significant differences, in terms of plant species diversity, composition, habitat condition and species attributes, between plant communities where non‐natives are present or abundant and those where non‐natives are absent or infrequent? 3. Are the observed differences generic to non‐natives or do individual non‐native species differ in their vegetation associations? Location: West Midlands Conurbation (WMC), UK. Methods: 56 sites were located randomly on four rivers across the WMC. Ten 2 m × 2 m quadrats were placed within 15 m of the river to sample vegetation within the floodplain at each site. All vascular plants were recorded along with site information such as surrounding land use and habitat types. Results: Non‐native species were found in many vegetation types and on all rivers in the WMC. There were higher numbers of non‐natives on more degraded, human‐modified rivers. More non‐native species were found in woodland, scrub and tall herb habitats than in grasslands. We distinguish two types of communities with non‐natives. In communities colonized following disturbance, in comparison to quadrats containing no non‐native species, those with non‐natives had higher species diversity and more forbs, annuals and shortlived monocarpic perennials. Native species in quadrats containing non‐natives were characteristic of conditions of higher fertility and pH, had a larger specific leaf area and were less stress tolerant or competitive. In later successional communities dominated by particular non‐natives, native diversity declined with increasing cover of non‐natives. Associated native species were characteristic of low light conditions. Conclusions: Communities containing non‐natives can be associated with particular types of native species. Extrinsic factors (disturbance, eutrophication) affected both native and non‐native species. In disturbed riparian habitats the key determinant of diversity is dominance by competitive invasive species regardless of their native or non‐native origin.     

Dominant forest tree mycorrhizal type mediates understory plant invasions

Forest mycorrhizal type mediates nutrient dynamics, which in turn can influence forest community structure and processes. Using forest inventory data, we explored how dominant forest tree mycorrhizal type affects understory plant invasions with consideration of forest structure and soil properties. We found that arbuscular mycorrhizal (AM) dominant forests, which are characterised by thin forest floors and low soil C : N ratio, were invaded to a greater extent by non‐native invasive species than ectomycorrhizal (ECM) dominant forests. Understory native species cover and richness had no strong associations with AM tree dominance. We also found no difference in the mycorrhizal type composition of understory invaders between AM and ECM dominant forests. Our results indicate that dominant forest tree mycorrhizal type is closely linked with understory invasions. The increased invader abundance in AM dominant forests can further facilitate nutrient cycling, leading to the alteration of ecosystem structure and functions.     

Nutrient enrichment enhances hidden differences in phenotype to drive a cryptic plant invasion

Many mechanisms of invasive species success have been elucidated, but those driving cryptic invasions of non‐native genotypes remain least understood. In one of the most successful cryptic plant invasions in North America, we investigate the mechanisms underlying the displacement of native Phragmites australis by its Eurasian counterpart. Since invasive Phragmites’ populations have been especially prolific along eutrophic shorelines, we conducted a two‐year field experiment involving native and invasive genotypes that manipulated nutrient level and competitor identity (inter‐ and intra‐genotypic competition) to assess their relative importance in driving the loss of native Phragmites. Inter‐genotypic competition suppressed aboveground biomass of both native and invasive plants regardless of nutrient treatment (～ 27%), while nutrient addition disproportionately enhanced the aboveground biomass (by 67%) and lateral expansion (by > 3 × farther) of invasive Phragmites. Excavation of experimental plots indicated that nutrient addition generates these differences in aboveground growth by differentially affecting rhizome production in invasive vs native plants; invasive rhizome biomass and rhizome length increased by 595% and 32% with nutrient addition, respectively, while natives increased by only 278% and 15%. Regardless of nutrient level, native rhizomes produced twice as many roots compared to invasives, which field surveys revealed are heavily infected with mycorrhizal symbionts. These results suggest that native Phragmites competes well under nutrient‐limited conditions because its rhizomes are laden with nutrient‐harvesting roots and mycorrhizae. Invasive Phragmites’ vigorous aboveground response to nutrients and scarcity of lateral roots, in contrast, may reflect its historic distribution in eutrophic Eurasian wetlands and correspond to its prevalence in New England marshes characterized by elevated nutrient availability and relaxed nutrient competition. These findings reveal that discrete differences in phenotype can interact with anthropogenic modification of environmental conditions to help explain the success of cryptic invaders.     

Modeling ecological minimum requirements for distribution of greater sage‐grouse leks: implications for population connectivity across their western range,U.S.A

Greater sage‐grouse Centrocercus urophasianus (Bonaparte) currently occupy approximately half of their historical distribution across western North America. Sage‐grouse are a candidate for endangered species listing due to habitat and population fragmentation coupled with inadequate regulation to control development in critical areas. Conservation planning would benefit from accurate maps delineating required habitats and movement corridors. However, developing a species distribution model that incorporates the diversity of habitats used by sage‐grouse across their widespread distribution has statistical and logistical challenges. We first identified the ecological minimums limiting sage‐grouse, mapped similarity to the multivariate set of minimums, and delineated connectivity across a 920,000 km² region. We partitioned a Mahalanobis D² model of habitat use into k separate additive components each representing independent combinations of species–habitat relationships to identify the ecological minimums required by sage‐grouse. We constructed the model from abiotic, land cover, and anthropogenic variables measured at leks (breeding) and surrounding areas within 5 km. We evaluated model partitions using a random subset of leks and historic locations and selected D² (k = 10) for mapping a habitat similarity index (HSI). Finally, we delineated connectivity by converting the mapped HSI to a resistance surface. Sage‐grouse required sagebrush‐dominated landscapes containing minimal levels of human land use. Sage‐grouse used relatively arid regions characterized by shallow slopes, even terrain, and low amounts of forest, grassland, and agriculture in the surrounding landscape. Most populations were interconnected although several outlying populations were isolated because of distance or lack of habitat corridors for exchange. Land management agencies currently are revising land‐use plans and designating critical habitat to conserve sage‐grouse and avoid endangered species listing. Our results identifying attributes important for delineating habitats or modeling connectivity will facilitate conservation and management of landscapes important for supporting current and future sage‐grouse populations.     

N/P imbalance as a key driver for the invasion of oligotrophic dune systems by a woody legume

Oligotrophic ecosystems, previously considered to be more resilient to invasive plants, are now recognised to be highly vulnerable to invasions. In these systems, woody legumes show belowground ecosystem engineering characteristics that enable invasion, however, the underlying processes are not well understood. Using a Portuguese primary dune ecosystem as an oligotrophic model system, belowground biomass pools, turnover rates and stoichiometry of a native (Stauracanthus spectabilis) and an invasive legume (Acacia longifolia) were compared and related to changes in the foliage of the surrounding native (Corema album) vegetation. We hypothesized that the invasive legume requires less phosphorus per unit of biomass produced and exhibits an enhanced nutrient turnover compared to the native vegetation, which could drive invasion by inducing a systemic N/P imbalance. Compared with the native legumes, A. longifolia plants had larger canopies, higher SOM levels and lower tissue P concentrations. These attributes were strongly related to legume influence as measured by increased foliar N content and less depleted δ¹⁵N signatures in the surrounding C. album vegetation. Furthermore, higher root N concentration and increased nutrient turnover in the rhizosphere of the invader were associated with depleted foliar P in C. album. Our results emphasize that while A. longifolia itself maintains an efficient phosphorus use in biomass production, at the same time it exerts a strong impact on the N/P balance of the native system. Moreover, this study highlights the engineering of a belowground structure of roots and rhizosphere as a crucial driver for invasion, due to its central role in nutrient turnover. These findings provide new evidence that, under nutrient‐limited conditions, considering co‐limitation and nutrient cycling in oligotrophic systems is essential to understand the engineering character of invasive woody legumes.     

Landscape characteristics of non‐native pine plantations and invasions in Southern Chile

The spread of non‐native conifers into areas naturally dominated by other vegetation types is a growing problem in South America. This process results in a landscape transformation as the conifers suppress native vegetation leading to reduced biodiversity, lower water availability and altered nutrient dynamics. Previous research highlights the broad spatial extents of land cover change in parts of Chile. However, in Southern Chile, the extent of plantations and the landscape characteristics associated with plantations and ongoing pine invasions are poorly understood. Here, we characterised non‐native pine land cover within one Landsat scene (World Reference System 2 Path 232/Row 92; ~34 000 km²) in Southern Chile. We created training data based on historical high‐resolution imagery, derived land cover predictors from time series of Landsat observations and used a Random Forest classifier to map the distribution of non‐native pines. The overall classification accuracy was 88%, and the accuracy of the non‐native pine class exceeded 90%. Although 71% of non‐native pine patches were within 500 m of other non‐native pine patches, isolated non‐native pine patches were found to occur up to 55 km from the nearest neighbour. These distant plantations could exacerbate invasion risk by creating propagule sources for novel invasion fronts. In relation to landscape characteristics, non‐native pines were found to be more likely to occur in low slope and mid‐elevation areas. Because most of the study area is native forest, most non‐native pine patches border native forest. However, non‐native pine patches were almost three times more likely than random patches to border grass/agriculture. This suggests that grasslands and disturbed sites, which have low resistance to non‐native pine invasion, are disproportionately exposed to pine propagules. Our results indicate that non‐native pine plantations are extensive across Southern Chile, and well poised to cause future invasion.     

Stable isotope and molecular analyses indicate that hybridization with non‐native domesticated common carp influence habitat use of native carp

Hybridization between native and non‐native species has consequences for survival and growth rates of hybrid offspring, but the influences on their functional roles such as habitat use are little studied and poorly understood. The Japanese native common carp Cyprinus carpio coexist and hybridize with non‐native domesticated carp in natural Japanese lakes. We have combined stable isotope and molecular information to examine whether habitat use of carp varies depending on the degree of hybridization between native and non‐native carp. We sampled 69 carp from Lake Kasumigaura where hybrid swarms between native and non‐native carp are advancing, evaluated the degree of hybridization for each individual by genotyping five single nucleotide polymorphism (SNP) markers, and analyzed their carbon and nitrogen stable isotopes. Although we did not find any genetically pure native carp in the lake, the results showed that carp δ¹³C increased with increasing frequency of non‐native alleles but that δ¹⁵N did not change. This indicates that non‐native carp use the littoral zone more frequently than native carp. This difference in habitat use was supported by a multisource mixing model, showing that the contribution of limnetic primary consumers to the diets of non‐native carp was lower than that of individuals with the highest frequency of native alleles. By combining two very different methods, our results thus suggest that multiple‐generation hybridization can influence habitat and resource use. Habitat partitioning should be considered when evaluating the genetic impacts of invasive species and races on native species and ecosystem processes.     

Modelling the distribution of a threatened habitat: the California sage scrub

Aim Using predictive species distribution and ecological niche modelling our objectives are: (1) to identify important climatic drivers of distribution at regional scales of a locally complex and dynamic system – California sage scrub; (2) to map suitable sage scrub habitat in California; and (3) to distinguish between bioclimatic niches of floristic groups within sage scrub to assess the conservation significance of analysing such species groups. Location Coastal mediterranean‐type shrublands of southern and central California. Methods Using point localities from georeferenced herbarium records, we modelled the potential distribution and bioclimatic envelopes of 14 characteristic sage scrub species and three floristic groups (south‐coastal, coastal–interior disjunct and broadly distributed species) based upon current climate conditions. Maxent was used to map climatically suitable habitat, while principal components analysis followed by canonical discriminant analysis were used to distinguish between floristic groups and visualize species and group distributions in multivariate ecological space. Results Geographical distribution patterns of individual species were mirrored in the habitat suitability maps of floristic groups, notably the disjunct distribution of the coastal–interior species. Overlap in the distributions of floristic groups was evident in both geographical and multivariate niche space; however, discriminant analysis confirmed the separability of floristic groups based on bioclimatic variables. Higher performance of floristic group models compared with sage scrub as a whole suggests that groups have differing climate requirements for habitat suitability at regional scales and that breaking sage scrub into floristic groups improves the discrimination between climatically suitable and unsuitable habitat. Main conclusions The finding that presence‐only data and climatic variables can produce useful information on habitat suitability of California sage scrub species and floristic groups at a regional scale has important implications for ongoing efforts of habitat restoration for sage scrub. In addition, modelling at a group level provides important information about the differences in climatic niches within California sage scrub. Finally, the high performance of our floristic group models highlights the potential a community‐level modelling approach holds for investigating plant distribution patterns.     

Multi‐trophic impacts of an invasive aquatic plant

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Effects of habitat transitions on rainforest bird communities across an anthropogenic landscape mosaic

We compared bird community responses to the habitat transitions of rainforest‐to‐pasture conversion, consequent habitat fragmentation, and post‐agricultural regeneration, across a landscape mosaic of about 600 km² in the eastern Australian subtropics. Birds were surveyed in seven habitats: continuous mature rainforest; two size classes of mature rainforest fragment (4–21 ha and 1–3 ha); regrowth forest patches dominated by a non‐native tree (2–20 ha, 30–50 years old); two types of isolated mature trees in pasture; and treeless pasture, with six sites per habitat. We compared the avifauna among habitats and among sites, at the levels of species, functional guilds, and community‐wide. Community‐wide species richness and abundance of birds in pasture sites were about one‐fifth and one‐third, respectively, of their values in mature rainforest (irrespective of patch size). Many measured attributes changed progressively across a gradient of increased habitat simplification. Rainforest specialists became less common and less diverse with decreased habitat patch size and vegetation maturity. However, even rainforest fragments of 1–3 ha supported about half of these species. Forest generalist species were largely insensitive to patch size and successional stage. Few species reached their greatest abundance in either small rainforest fragments or regrowth. All pastures were dominated by bird species whose typical native habitats were grassland, wetland, and open eucalypt forest, while pasture trees modestly enhanced local bird communities. Overall, even small scattered patches of mature and regrowth forest contributed substantial bird diversity to local landscapes. Therefore, maximizing the aggregate rainforest area is a useful regional conservation strategy.     

The Potential Role of Stream Habitat Restoration in Facilitating Salmonid Invasions: A Habitat‐Hydraulic Modeling Approach

Many successful invasions have taken place in systems where harmful disturbance has changed habitat conditions. However, less attention has been paid to the role of habitat restoration, which modifies habitats and thus also has the potential to facilitate invasions. We examined whether in‐stream habitat restorations have the potential to either facilitate or resist invasion by two widely introduced non‐native stream salmonids, Salvelinus fontinalis Mitchill and Oncorhynchus mykiss Walbaum, in Finland. A physical habitat simulation system was used to calculate whether the habitat area for the target species increased or decreased following the restorations. For comparison, we also reported results for four native stream fish species. The simulations showed that the restored streams provided the highest amount of usable habitat area for the native species, particularly for Salmo salar L. and Gottus gobio L. However, it was interesting to note that the restorations significantly increased habitat quality for the two non‐native species, especially at low flows. Nevertheless, the non‐native species had the lowest amount of usable habitat area overall. The modeling results indicated that not only habitat destruction but also habitat restoration could contribute to the spread of non‐native species. Fisheries and wildlife managers should be aware of the possibility, when restoring habitats in order to preserve native ecosystems, that non‐native species could manage to gain a foothold in restored habitats and use them as population sources for further spread. Knowing the widespread negative effect of non‐native species, this risk should not be underestimated.     

Occurrence of non‐native fishes in the Danube east of Vienna (Austria) and potential interactions of invasive gobiids with native fishes

Non‐native fish species pose a major threat to local fish populations and aquatic ecosystems in general. Invasive gobies are a particular focus of research, but with partly inconsistent results. While some studies reported severe detrimental impacts on native species, others have concluded less serious or neutral effects. We provide results from a large‐scale, multi‐annual fish monitoring program on the occurrence and abundance of non‐native fishes in the main stem of a free‐flowing section of the Austrian Danube. Special emphasis was placed on identifying positive or negative interactions of invasive gobies with native species. Whereas most non‐native species occurred too sporadically or were too few in number to infer a direct threat on the local fish community, invasive gobies were among the most common fishes throughout all sampling years. Co‐occurrence analyses revealed species‐ and mesohabitat type‐specific associations of gobies with native species, which were primarily positive. Notably, native predators such as asp, burbot, or perch probably benefit from the ubiquitous gobies. Two characteristic fluvial fishes revealed negative associations with invasive gobies, namely barbel (Barbus barbus) and Danube whitefin gudgeon (Romanogobio vladykovi): they appear to avoid habitats occupied by gobies. Accordingly, high abundances of round and bighead goby most likely resulted in population losses of barbel and whitefin gudgeon, respectively. Overall, our results indicate a limited negative impact of non‐native species in the sampling area. This is because only two out of 51 occurring species were found to be adversely affected by gobies, the share of co‐occurrences with native species was high, and other non‐native species were generally rare. Nevertheless, invasions are highly dynamic, and new non‐native species are likely to occur in the Austrian Danube, calling for continued monitoring and awareness.