Widespread hybridization between native Atlantic salmon, Salmo salar, and introduced brown trout, S. trutta, in eastern Newfoundland

Hybridization between native Atlantic salmon and introduced brown trout was found to occur at a mean frequency of 0.9% in Atlantic salmon populations in eastern Newfoundland. Hybrids were detected in five of the 10 watersheds studied, but consideration of sampling error suggests that they could have been present in the remaining five watersheds although they were not detected. The frequency found in the Newfoundland and other North American salmon populations is significantly greater than the 0.3% reported for salmon populations in the British Isles, and both are higher than frequencies observed in salmon populations in Sweden. The higher frequency in North America is in accord with the prediction that hybridization between species will be more frequent where one species is introduced than in areas where both are native.     

Spatial and temporal variability in propagule limitation of California native grasses

Community composition and diversity arise from limitations in propagule supply (i.e. propagule or seed limitation) and propagule establishment after arriving at a site (i.e. establishment or microsite limitation). Recent meta‐analyses suggest that the degree of propagule limitation depends on local abiotic and biotic conditions, which in turn are likely to vary spatially and temporally. Nevertheless, seed addition studies testing propagule limitation are rarely replicated in multiple locations and years and often lack experimental manipulations of critical determinants of propagule limitation, such as the density and species richness of the propagule pool and disturbance and resource supply of recipient site. The invasion of California (USA) grasslands by exotic annual species from the Mediterranean region is unique in its scope (over 9 million ha) and persistence (more than 150 years). This invasion provides an exciting context in which to test the role of spatial and temporal variability in mediating propagule limitation and ultimately the potential for restoration. Here I present the results of native‐grass, seed‐addition trials conducted along a 500 km gradient in California in three consecutive years spanning a wide range of environmental conditions and initial conditions (e.g. rainfall, seeding density and disturbance). While native grasses were able to establish at many locations, per capita seed survival was low suggesting that the fate of these species is governed by interplay between propagule and establishment limitation. Establishment primarily varied spatially, with the most successful establishment occurring at fertile locations with low resident species richness. While recruitment was highly variable among years initially, there was no difference among seedling trials after three years.     

Re‐building brown trout populations in dredged boreal forest streams: in‐stream restoration combined with stocking of young trout

1. Rivers in boreal forested areas were often dredged to facilitate the transport of timber resulting in channels with simplified bed structure and flow fields and reduced habitat suitability for stream organisms, especially lotic fishes. Currently, many streams are being restored to improve their physical habitat, by replacing boulders and gravel and removing constraining embankments. The most compelling justification behind stream restoration of former floatways has been the enhancement of native fish populations, specifically salmonids. 2. We examined the success of a stream management programme aimed at re‐building diminished brown trout (Salmo trutta) populations by monitoring densities of young‐of‐year and older trout in 18 managed and three reference streams during 2000–2005. Rehabilitation included in‐stream restoration combined with a 5‐year post‐restoration period of stocking young brown trout. Our space‐for‐time substitution design comprised four pre‐management, four under‐management, 10 post‐management and three reference streams. 3. Densities of young‐of‐year brown trout, indicating population establishment, were significantly higher in post‐ compared with pre‐management streams. However, density of young‐of‐year brown trout in post‐management streams was significantly lower compared with near‐pristine reference streams. Furthermore, success of managed brown trout population re‐building varied, indicating stream‐specific responses to management measures. Density of burbot (Lota lota), a native generalist predator, was associated with low recruitment of brown trout. 4. Stream‐specific responses imply that rehabilitation of brown trout populations cannot be precisely predicted thereby limiting application. Our findings support the importance of adaptive stream restoration and management, with focus on identifying factor(s) limiting the establishment of target fish populations.     

The abiotic and biotic factors limiting establishment of predatory fishes at their expanding northern range boundaries in Ontario,Canada

There is a poor understanding of the importance of biotic interactions in determining species distributions with climate change. Theory from invasion biology suggests that the success of species introductions outside of their historical ranges may be either positively (biotic acceptance) or negatively (biotic resistance) related to native biodiversity. Using data on fish community composition from two survey periods separated by approximately 28 years during which climate was warming, we examined the factors influencing the establishment of three predatory centrarchids: Smallmouth Bass (Micropterus dolomieu), Largemouth Bass (M. salmoides), and Rock Bass (Ambloplites rupestris) in lakes at their expanding northern range boundaries in Ontario. Variance partitioning demonstrated that, at a regional scale, abiotic factors play a stronger role in determining the establishment of these species than biotic factors. Pairing lakes within watersheds where each species had established with lakes sharing similar abiotic conditions where the species had not established revealed both positive and negative relationships between the establishment of centrarchids and the historical presence of other predatory species. The establishment of these species near their northern range boundaries is primarily determined by abiotic factors at a regional scale; however, biotic factors become important at the lake‐to‐lake scale. Studies of exotic species invasions have previously highlighted how spatial scale mediates the importance of abiotic vs. biotic factors on species establishment. Our study demonstrates how concepts from invasion biology can inform our understanding of the factors controlling species distributions with changing climate.     

Propagule pressure governs establishment of an invasive herb

The success of plant invasions may be limited by the availability of propagules and/or of suitable microsites, with microsite availability being affected by, for example, disturbance and interspecific competition. A mechanistic understanding of the contributions of propagule pressure and microsite limitation to plant invasions is therefore required to minimise future invasions. Here, we investigated the relative roles of propagule pressure, the availability of microsites, and their interaction on the establishment of an invasive herb, Lupinus polyphyllus, in two geographic regions representing different climate and growth conditions in Finland (a more productive southern region and a harsher central region). We carried out a field experiment in 14 L. polyphyllus populations, in which we manipulated both propagule pressure and disturbance. In a complementary greenhouse experiment, we manipulated propagule pressure and interspecific competition. Seedling establishment of L. polyphyllus was higher in the more productive southern region than in the harsher central region. The number of L. polyphyllus seedlings increased with increasing propagule pressure regardless of disturbance or interspecific competition. However, the number of L. polyphyllus seedlings per sown seed (relative establishment) tended to decrease with increasing propagule pressure, indicating that the positive effect of propagule pressure on early invasion is partially counteracted by density-dependent mortality at high seed densities. Our results highlight the dominant role of propagule pressure over disturbance and interspecific competition in the establishment of L. polyphyllus, suggesting that the early stage of invasion is limited by the availability of propagules rather than the availability of suitable microsites.     

Ecological resistance, seed density and their interactions determine patterns of invasion in a California coastal grassland

Relatively little experimental evidence is available regarding how ecological resistance and propagule density interact in their effects on the establishment of invasive exotic species. We examined the independent and interactive effects of neighbour cover (biotic resistance), winter vs. spring water addition (abiotic resistance) and seed density on the invasion of the European perennial grass Holcus lanatus into a California coastal grassland dominated by exotic annual grasses. We found that decreased competition from resident exotic grasses had no effect. In contrast, increased late-season water availability eroded the abiotic resistance offered by naturally dry conditions, facilitating invasion. Finally, watering treatment and seed density interacted strongly in determining seedling survival: while seedling mortality was close to 100% in ambient and winter water addition plots, survivor numbers increased with seed density in spring-watered plots. Thus, decreased abiotic resistance can amplify the effect of increased propagule density on seedling establishment, thereby increasing the likelihood of invasion.     

Genetic analysis of a novel invasion of Puerto Rico by an exotic constricting snake

The tropical island Puerto Rico is potentially vulnerable to invasion by some species of exotic snakes; however, until now no established populations had been reported. Here we report and genetically characterize the nascent invasion of Puerto Rico by an exotic constricting snake of the family Boidae (Boa constrictor) using mtDNA and microsatellite data. Over 150 individual B. constrictor have been removed from Mayagüez municipality since May 2011, and our results from the genetic analysis of 32 individuals suggest that this population was recently founded by individuals of one subspecies from a genetic lineage common to zoo and breeding collections, but that the potential propagule pool consists of two subspecies. We also suggest that anthropogenic long-distance dispersal within the island of Puerto Rico may be occurring from the established population, with implications for further establishment across the island. This study represents the first report of the naturalization of an invasive species of boid snake in Puerto Rico and will be important in determining mitigation strategies for this invasion as well as providing a basis for comparison to other on-going studies of invasive snakes.     

Culvert Replacement and Stream Habitat Restoration: Implications from Brook Trout Management in an Appalachian Watershed, U.S.A.

Large-scale culvert replacement programs could benefit migratory fish populations by reconnecting reproductive and foraging habitats in fragmented watersheds. The objectives of this study were to: (1) identify stream and culvert characteristics contributing to fish passage barriers within an Appalachian watershed, U.S.A.; (2) quantify the total amount of Brook trout ( Salvelinus fontinalis ) reproductive habitat isolated above culverts; and (3) use an ecological currency to identify culvert replacement priorities and stream mitigation credit opportunities. We surveyed 120 state-owned culverts and used a fish passage assessment filter to determine the "passability" of each culvert. We then constructed a geographic information system stream network model to quantify the amount of trout reproductive habitat isolated by culverts. Ninety-seven percent of surveyed culverts were classified as obstacles or complete barriers to trout dispersal. Culvert impassability was higher in small streams with slopes exceeding 3–5%, suggesting a direct relationship between slope and impassability. Thirty-three percent of Brook trout reproductive habitat, representing over 200 km of stream, was isolated by culverts. This is a conservative estimate, because we did not survey privately or federally owned culverts. The top 20 prioritized culverts accounted for nearly half of the habitat loss. Our results indicate that standard culvert designs placed in streams with slopes exceeding 5% consistently produce trout dispersal barriers and should be avoided during new road construction. The process developed here provides an efficient method for identifying culvert replacement priorities and may be used to maximize watershed scale benefits of stream restoration.     

Interspecific association of brown trout (Salmo trutta) with non‐native brook trout (Salvelinus fontinalis) at the fry stage

The introduction of non‐native brook trout (Salvelinus fontinalis) in Europe has led to displacement and decreasing populations of native brown trout (Salmo trutta). Some studies have found that brown trout shift to a diet niche similar to brook trout when the two species live in sympatry, which conflicts with the competitive exclusion principle. A change in feeding niche may be a sign of early interspecific association and social learning, leading to behavioral changes. As a first step to address this possibility, it is essential to assess the interspecific association between the species during the early ontogenetic life stages. In this study, we therefore assess whether juvenile brown trout associate with non‐native juvenile brook trout to the same extent as with conspecifics by setting up two experiments: (i) a binomial choice test allowing visual and chemical cues to estimate the species specificity of group preference, and (ii) an association test without physical barriers to estimate the degree of association of a focal brown trout with a group of either conspecifics or heterospecifics. In experiment (1), we found that focal juvenile brown trout preferred to associate with the stimuli groups and did not discriminate either against conspecific or heterospecific groups. Furthermore, more active individuals showed stronger preference for the stimuli group than less active ones, regardless of species. In experiment (2), we found that brook trout groups had a tighter group structure than brown trout groups, and that focal brown trout showed stronger association with brook trout than with brown trout. These results indicate that brown trout may associate with brook trout at an early life stage, which would allow for interspecific social learning to occur. Future studies should look closer into causes and consequences of interspecific association and social learning, including potential effects on the phenotype selection in brown trout populations.     

Temporal and nonlinear dispersal patterns of <Emphasis Type="Italic">Ludwigia hexapetala</Emphasis> in a regulated river

Rivers are vulnerable to biological invasion due to hydrologic connectivity, which facilitates post-entry movement of aquatic plant propagules by water currents. Ecological and watershed factors may influence spatial and temporal dispersal patterns. Field-based data on dispersal could improve risk assessment models and management responses. Ludwigia hexapetala, an invasive emergent macrophyte, provides a case study for understanding dispersal patterns throughout a watershed. The species spreads via hydrochory and is increasingly imposing detrimental ecological and economic impacts within watersheds of the United States and Europe. We investigated morphology of shoot fragments and their dispersal in the Russian River watershed of California, capturing shoot fragments of L. hexapetala during repeated summer surveys at five locations in the river and quantifying their morphological traits that predict establishment success. Highly variable capture counts suggest the importance of pulse disturbance events in local dispersal of L. hexapetala. Unexpectedly, dispersing propagule pressure was nonlinear, with more shoot fragments captured in the middle rather than lower river. Captured fragments in the middle river were twice the length of fragments captured in the lower river and bore 83% more stem nodes, characteristics associated with greater establishment success. Our results support development of spatially targeted management, outreach, and prevention efforts that could lead to decreased propagule pressure in the watershed.     

Propagule pressure and native community connectivity interact to influence invasion success in metacommunities

Mechanistic insights from invasion biology indicate that propagule pressure of exotic species and native community structure can independently influence establishment success. The role of native community connectivity via species dispersal and its potential interaction with propagule pressure on invasion success in metacommunities, however, remains unknown. Native community connectivity may increase biotic resistance to invasion by enhancing species richness and evenness, but the effects could depend upon the level of propagule pressure. In this study, a mesocosm experiment was used to evaluate the independent and combined effects of exotic propagule pressure and native community connectivity on invasion success. The effects of three levels of exotic Daphnia lumholtzi propagule pressure on establishment success, community structure and ecosystem attributes were evaluated in native zooplankton communities connected by species dispersal versus unconnected communities, and relative to a control without native species. Establishment of the exotic species exhibited a propagule dose‐dependent relationship with high levels of propagule pressure resulting in the greatest establishment success. Native community connectivity, however, effectively reduced establishment at the low level of propagule pressure and further augmented native species richness across propagule pressure treatments. Propagule pressure largely determined the negative impacts of the exotic species on native species richness, native biomass and edible producer biomass. The results highlight that native community connectivity can reduce invasion success at a low propagule dose and decrease extinction risk of native competitors, but high propagule pressure can overcome connectivity‐mediated biotic resistance to influence establishment and impact of the exotic species. Together, the results emphasize the importance of the interaction of propagule pressure and community connectivity as a regulator of invasion success, and argue for the maintenance of metacommunity connectivity to confer invasion resistance.     

Watershed‐level brook trout genetic structuring: Evaluation and application of riverscape genetics models

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The importance of disturbance by fire and other abiotic and biotic factors in driving cheatgrass invasion varies based on invasion stage

Disturbances create fluctuations in resource availability that alter abiotic and biotic constraints. Exotic invader response may be due to multiple factors related to disturbance regimes and complex interactions between other small- and large-scale abiotic and biotic processes that may vary across invasion stages. We explore how cheatgrass responds to both frequency and season of prescribed burning for a 10-year period in ponderosa pine forested stands. To understand interactions of fire disturbance, other abiotic factors, biotic resistance, and propagule pressure, we use long-term data from different spatial scales representing different invasion stages (local establishment or spread and broader scale extent/impact) to model cheatgrass dynamics. We found that after 10 years, cheatgrass cover increased with fall burning regardless of burn frequency (1 burn vs. 3 burns). There was no evidence that cheatgrass invasion is decreasing through time even in areas burned only once. Factors important for explaining local fine-scale cheatgrass establishment and spread, and broader scale extent/impact varied. The spatial extent of the first burns facilitated fine-scale cheatgrass establishment while bare soil cover constrained establishment. Biotic resistance, in the form of native annual forb cover, constrained fine-scale cheatgrass spread. Initial cheatgrass abundance in 2002, a factor related to propagule pressure, was key for explaining the broader scale extent/impact of cheatgrass by 2012. Biotic resistance, in the form of native perennial grass cover, constrained extent/impact but only when initial cheatgrass abundance was low. Our findings regarding factors affecting invasion dynamics may be useful to consider for future restoration and conservation efforts in burned ponderosa pine forests.     

Genetic diversity is related to climatic variation and vulnerability in threatened bull trout

Understanding how climatic variation influences ecological and evolutionary processes is crucial for informed conservation decision‐making. Nevertheless, few studies have measured how climatic variation influences genetic diversity within populations or how genetic diversity is distributed across space relative to future climatic stress. Here, we tested whether patterns of genetic diversity (allelic richness) were related to climatic variation and habitat features in 130 bull trout (Salvelinus confluentus) populations from 24 watersheds (i.e., ~4–7th order river subbasins) across the Columbia River Basin, USA. We then determined whether bull trout genetic diversity was related to climate vulnerability at the watershed scale, which we quantified on the basis of exposure to future climatic conditions (projected scenarios for the 2040s) and existing habitat complexity. We found a strong gradient in genetic diversity in bull trout populations across the Columbia River Basin, where populations located in the most upstream headwater areas had the greatest genetic diversity. After accounting for spatial patterns with linear mixed models, allelic richness in bull trout populations was positively related to habitat patch size and complexity, and negatively related to maximum summer temperature and the frequency of winter flooding. These relationships strongly suggest that climatic variation influences evolutionary processes in this threatened species and that genetic diversity will likely decrease due to future climate change. Vulnerability at a watershed scale was negatively correlated with average genetic diversity (r = ?0.77; P < 0.001); watersheds containing populations with lower average genetic diversity generally had the lowest habitat complexity, warmest stream temperatures, and greatest frequency of winter flooding. Together, these findings have important conservation implications for bull trout and other imperiled species. Genetic diversity is already depressed where climatic vulnerability is highest; it will likely erode further in the very places where diversity may be most needed for future persistence.     

Consequences of climatic change for water temperature and brown trout populations in Alpine rivers and streams

Twenty‐five years of extensive water temperature data show regionally coherent warming to have occurred in Alpine rivers and streams at all altitudes, reflecting changes in regional air temperature. Much of this warming occurred abruptly in 1987/1988. For brown trout populations, the warming resulted in an upward shift in thermal habitat that was accelerated by an increase in the incidence of temperature‐dependent Proliferative Kidney Disease at the habitat's lower boundary. Because physical barriers restrict longitudinal migration in mountain regions, an upward habitat shift in effect implies habitat reduction, suggesting the likelihood of an overall population decrease. Extensive brown trout catch data documenting an altitudinally dependent decline indicate that such a climate‐related population decrease has in fact occurred. Our analysis employs a quantitatively defined reference optimum temperature range for brown trout, based on the sinusoidal regression of seasonally varying field data.     

Does restoration work? It depends on how we measure success

The restoration of 4 partial stream barriers was evaluated in watersheds of Terra Nova National Park, Newfoundland, Canada from 2009 to 2011. Brook trout (n = 462) were tagged and tracked moving through our study sites using passive‐integrated transponder telemetry and the restoration actions were assessed using 3 different measures: passage success rates; the range of passable flows; and the availability of passable flows. We considered the observed results within a before‐after‐control‐impact (BACI) design that included reference reaches and pre‐restoration observations. The conclusions of BACI analyses were also contrasted with those that would have been obtained from commonly used before‐after (B‐A) or control‐impact (C‐I) study designs. While the restoration actions changed hydrological conditions in a way that should facilitate fish passage, our biological measures indicated that success was variable across culverts and within culverts depending on the measure evaluated. Furthermore, the natural temporal and spatial variability of fish movements often resulted in different conclusions between the more robust BACI design and the more commonly used B‐A and C‐I designs. Our results demonstrate that restoration of partial barriers may not always yield dramatic improvements. Furthermore, without suitable controls, the chances of drawing false conclusions regarding restorations in temporally and spatially dynamic systems are substantial.     

Effects of stocking on the genetic diversity of brown trout populations of the Adriatic and Danubian drainages in Switzerland

This study focuses on genetic variation of brown trout Salmo trutta populations of the Adriatic and Danubian drainages in Switzerland. The allozyme and other protein loci data show a major replacement of native stocks from the Adriatic drainages by introduced hatchery trout of Atlantic basin origin. In most samples, diagnostic alleles for the Adriatic form of Salmo trutta f. fario and for the marbled trout Salmo trutta marmoratus are found at very low frequencies (f<0.15). Taking into account previous genetic studies on brown trout of this basin, the Danubian samples are not heavily contaminated with foreign alleles. The results are consistent with records of local stocking activities which account in part for the high introgression rates of Atlantic alleles into local populations of the Adriatic drainages. In addition, introgression is enhanced by a decrease of natural reproduction which is caused by a deterioration of trout habitats through human activities. Furthermore, a third mechanism is proposed that may contribute to the high introgression rates observed: if Atlantic trout are introduced, the reproductive barriers between the two native forms, marbled trout and Adriatic fario respectively, break down. Atlantic trout apparently hybridize with both native forms and generate gene flow between them. In some parts of Adriatic drainages in Switzerland, the patterns of introgression and hybridization are further complicated by introduction of trout from the Danubian system. Alleles of the marbled trout are also found in the samples of the Danubian drainage system. These are due to stocking activities across the watershed.     

Non-native species limit stream restoration benefits for brook trout

Success of stream restoration can be difficult to define because many interacting abiotic and biotic factors across spatio-temporal scales can have measurable effects. Consequently, failure in habitat restoration to achieve targeted biological goals may reflect interactions of habitat restoration with unaccounted risks that have yet to be addressed on the landscape. This is particularly true within invaded landscapes, where habitat restoration can benefit non-native competitors as much as the native fishes for which restoration is designed. We tested for interacting effects of a reach scale habitat restoration effort and non-native trout competition on habitat use by a brook trout (Salvelinus fontinalis) metapopulation within a productive main stem corridor of the Shavers Fork watershed, West Virginia. We used a joint species occupancy model within a BACI sampling design to show that brook trout occupancy of main stem habitat was highest post-restoration within restored sampling reaches, but this benefit to native brook trout was conditional on brown trout (Salmo trutta) not being present within the main stem habitat. Collectively these results indicate that habitat restoration was only beneficial for native brook trout when non-native trout were absent from the restored sampling area. Proactive approaches to restoration will be integral for supporting resilient ecosystems in response to future anthropogenic threats (e.g. climate change), and we have shown that such actions will only be successful if non-native competitors do not also benefit from the restoration actions.     

Human-related processes drive the richness of exotic birds in Europe

Both human-related and natural factors can affect the establishment and distribution of exotic species. Understanding the relative role of the different factors has important scientific and applied implications. Here, we examined the relative effect of human-related and natural factors in determining the richness of exotic bird species established across Europe. Using hierarchical partitioning, which controls for covariation among factors, we show that the most important factor is the human-related community-level propagule pressure (the number of exotic species introduced), which is often not included in invasion studies due to the lack of information for this early stage in the invasion process. Another, though less important, factor was the human footprint (an index that includes human population size, land use and infrastructure). Biotic and abiotic factors of the environment were of minor importance in shaping the number of established birds when tested at a European extent using 50 x 50 km2 grid squares. We provide, to our knowledge, the first map of the distribution of exotic bird richness in Europe. The richest hotspot of established exotic birds is located in southeastern England, followed by areas in Belgium and The Netherlands. Community-level propagule pressure remains the major factor shaping the distribution of exotic birds also when tested for the UK separately. Thus, studies examining the patterns of establishment should aim at collecting the crucial and hard-to-find information on community-level propagule pressure or develop reliable surrogates for estimating this factor. Allowing future introductions of exotic birds into Europe should be reconsidered carefully, as the number of introduced species is basically the main factor that determines the number established.