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Post-fire recovery of sagebrush-steppe communities is better explained by elevation than climate-derived indicators of resistance and resilience

More landscapes require restoration than can feasibly be treated, and so decision-support tools to prioritize areas for treatment are needed. Moreover, restoration is complicated by the threat of biological invasion in disturbed areas, and so indicators of ecosystem resistance to invasion and resilience to disturbance (hereafter R&R) are important candidate criteria for prioritizing sites for restoration. We asked how climate-based R&R indicators that differed in being either categorical or continuous compared in their ability to explain plant-community recovery after six wildfires that collectively encompassed >750,000 ha and 7803 plot-year observations in sagebrush steppe of the western USA. Unique associations of species that most frequently co-occurred were identified using structural topic modelling. Mixed effect random forests were used to identify the relative importance of various R&R indicators in explaining post-fire plant associations compared with weather, landscape characteristics and treatment history. Simple metrics (elevation, latitude, longitude and year of monitoring) were more informative predictors of post-fire recovery than climate-based R&R indicators. However, small differences in the abundances of perennial grass and especially annual grass associations were predicted by the spring modified Thornthwaite Moisture Index (difference between precipitation and potential evapotranspiration). Synthesis and applications : The convenience of categorical resistance and resilience indicators has led to their widespread adoption for large-scale planning of restoration. Our results reveal that none of the resistance and resilience indicators assessed effectively explained post-fire restoration better than elevation, although a simple continuous resistance and resilience indicator describing water balance performed better than categorical indicators for explaining small but critical differences in cheatgrass association abundances.

California, Idaho, Nevada, Oregon

A framework to integrate innovations in invasion science for proactive management

Invasive alien species (IAS) are a rising threat to biodiversity, national security, and regional economies, with impacts in the hundreds of billions of U.S. dollars annually. Proactive or predictive approaches guided by scientific knowledge are essential to keeping pace with growing impacts of invasions under climate change. Although the rapid development of diverse technologies and approaches has produced tools with the potential to greatly accelerate invasion research and management, innovation has far outpaced implementation and coordination. Technological and methodological syntheses are urgently needed to close the growing implementation gap and facilitate interdisciplinary collaboration and synergy among evolving disciplines. A broad review is necessary to demonstrate the utility and relevance of work in diverse fields to generate actionable science for the ongoing invasion crisis. Here, we review such advances in relevant fields including remote sensing, epidemiology, big data analytics, environmental DNA (eDNA) sampling, genomics, and others, and present a generalized framework for distilling existing and emerging data into products for proactive IAS research and management. This integrated workflow provides a pathway for scientists and practitioners in diverse disciplines to contribute to applied invasion biology in a coordinated, synergistic, and scalable manner.

Biological Reviews

Global phylogeography of the avian malaria pathogen Plasmodium relictum based on MSP1 allelic diversity

Knowing the genetic variation that occurs in pathogen populations and how it is distributed across geographical areas is essential to understand parasite epidemiology, local patterns of virulence, and evolution of host-resistance. In addition, it is important to identify populations of pathogens that are evolutionarily independent and thus ‘free’ to adapt to hosts and environments. Here, we investigated genetic variation in the globally distributed, highly invasive avian malaria parasite Plasmodium relictum , which has several distinctive mitochondrial haplotyps (cyt b lineages, SGS1, GRW11 and GRW4). The phylogeography of P. relictum was accessed using the highly variable nuclear gene merozoite surface protein 1 (MSP1), a gene linked to the invasion biology of the parasite. We show that the lineage GRW4 is evolutionarily independent of GRW11 and SGS1 whereas GRW11 and SGS1 share MSP1 alleles and thus suggesting the presence of two distinct species (GRW4 versus SGS1 and GRW11). Further, there were significant differences in the global distribution of MSP1 alleles with differences between GRW4 alleles in the New and the Old World. For SGS1, a lineage formerly believed to have both tropical and temperate transmission, there were clear differences in MSP1 alleles transmitted in tropical Africa compared to the temperate regions of Europe and Asia. Further, we highlight the occurrence of multiple MSP1 alleles in GRW4 isolates from the Hawaiian Islands, where the parasite has contributed to declines and extinctions of endemic forest birds since it was introduced. This study stresses the importance of multiple independent loci for understanding patterns of transmission and evolutionary independence across avian malaria parasites.

Ecography

Catchment-scale determinants of nonindigenous minnow richness in the eastern United States

Understanding the drivers of biological invasions is critical for preserving aquatic biodiversity. Stream fishes make excellent model taxa for examining mechanisms driving species introduction success because their distributions are naturally limited by catchment boundaries. In this study, we compared the relative importance of catchment-scale abiotic and biotic predictors of native and nonindigenous minnow (Cyprinidae) richness in 170 catchments throughout the eastern United States. We compared historic and contemporary cyprinid distributional data to determine catchment-wise native/nonindigenous status for 152 species. Catchment-scale model predictor variables described natural (elevation, precipitation, flow accumulation) and anthropogenic (developed land cover, number of dams) abiotic features, as well as native congener richness. Native congener richness may represent either biotic resistance via interspecific competition, or trait preadaptation according to Darwin's naturalisation hypothesis. We used generalised linear mixed models to examine evidence supporting the relative roles of abiotic and biotic predictors of cyprinid introduction success. Native congener richness was positively correlated with nonindigenous cyprinid richness and was the most important variable predicting nonindigenous cyprinid richness. Mean elevation had a weak positive effect, and effects of other abiotic factors were insignificant and less important. Our results suggest that at this spatial scale, trait preadaptation may be more important than intrageneric competition for determining richness of nonindigenous fishes.

Ecology of Freshwater Fish

A trial of two trouts: Comparing the impacts of rainbow and brown trout on a native galaxiid

Rainbow trout Oncorhynchus mykiss and brown trout Salmo trutta are the world's two most widespread exotic fishes, dominate the fish communities of most cold-temperate waters in the southern hemisphere and are implicated in the decline and extirpation of native fish species. Here, we provide the first direct comparison of the impacts of rainbow and brown trout on populations of a native fish by quantifying three components of exotic species impact: range, abundance and effect. We surveyed 54 small streams on the island of Chilo?? in Chilean Patagonia and found that the rainbow trout has colonized significantly more streams and has a wider geographic range than brown trout. The two species had similar post-yearling abundances in allopatry and sympatry, and their abundances depended similarly on reach-level variation in the physical habitat. The species appeared to have dramatically different effects on native drift-feeding Aplochiton spp., which were virtually absent from streams invaded by brown trout but shared a broad sympatric range with rainbow trout. Within this range, the species' post-yearling abundances varied independently before and after controlling for variation in the physical habitat. In the north of the island, Aplochiton spp. inhabited streams uninvaded by exotic trouts. Our results provide a context for investigating the mechanisms responsible for apparent differences in rainbow and brown trout invasion biology and can help inform conservation strategies for native fishes in Chilo?? and elsewhere. ?? 2010 The Authors. Journal compilation ?? 2010 The Zoological Society of London.

Animal Conservation

Sonoran Desert ecosystem transformation by a C 4 grass without the grass/fire cycle

Aim Biological invasions facilitate ecosystem transformation by altering the structure and function, diversity, dominance and disturbance regimes. A classic case is the grass–fire cycle in which grass invasion increases the frequency, scale and/or intensity of wildfires and promotes the continued invasion of invasive grasses. Despite wide acceptance of the grass–fire cycle, questions linger about the relative roles that interspecific plant competition and fire play in ecosystem transformations. Location Sonoran Desert Arizona Upland of the Santa Catalina Mountains, Arizona, USA. Methods We measured species cover, density and saguaro ( Carnegiea gigantea ) size structure along gradients of Pennisetum ciliare invasion at 10 unburned/ungrazed P. ciliare patches. Regression models quantified differences in diversity, cover and density with respect to P. ciliare cover, and residence time and a Fisher's exact test detected demographic changes in saguaro populations. Because P. ciliare may have initially invaded locations that were both more invasible and less diverse, we ran analyses with and without the plots in which initial infestations were located. Results Richness and diversity decreased with P. ciliare cover as did cover and density of most dominant species. Richness and diversity declined with increasing time since invasion, suggesting an ongoing transformation. The proportion of old-to-young Carnegiea gigantea was significantly lower in plots with dominant P. ciliare cover. Main conclusions Rich desert scrub (15–25 species per plot) was transformed into depauperate grassland (2–5 species per plot) within 20 years following P. ciliare invasion without changes to the fire regime. While the onset of a grass–fire cycle may drive ecosystem change in the later stages and larger scales of grass invasions of arid lands, competition by P. ciliare can drive small-scale transformations earlier in the invasion. Linking competition-induced transformation rates with spatially explicit models of spread may be necessary for predicting landscape-level impacts on ecosystem processes in advance of a grass–fire cycle.

Arizona

Discordant introgression in a rapidly expanding hybrid swarm

The erosion of species boundaries can involve rapid evolutionary change. Consequently, many aspects of the process remain poorly understood, including the formation, expansion, and evolution of hybrid swarms. Biological invasions involving hybridization present exceptional opportunities to study the erosion of species boundaries because timelines of interactions and outcomes are frequently well known. Here, we examined clinal variation across codominant and maternally inherited genetic markers as well as phenotypic traits to characterize the expansion and evolution of a hybrid swarm between native Cyprinella venusta and invasive Cyprinella lutrensis minnows. Discordant introgression of phenotype, microsatellite multilocus genotype, and mtDNA haplotype indicates that the observable expansion of the C. venusta x C. lutrensis hybrid swarm is a false invasion front. Both parental and hybrid individuals closely resembling C. lutrensis are numerically dominant in the expansion wake, indicating that the non-native parental phenotype may be selectively favored. These findings show that cryptic introgression can extend beyond the phenotypic boundaries of hybrid swarms and that hybrid swarms likely expand more rapidly than can be documented from phenotypic variation alone. Similarly, dominance of a single parental phenotype following an introduction event may lead to instances of species erosion being mistaken for species displacement without hybridization.

Evolutionary Applications

Movement and behavioral states of common carp (Cyprinus carpio) in response to a behavioral deterrent in a navigational lock

Freshwater ecosystems are some of the most affected by biological invasions due, in part, to the introduction of invasive carp worldwide. Where carp have become established, management programs often seek to limit further range expansion into new areas by reducing their movement through interconnected rivers and waterways. Lock and dams are important locations for non-physical deterrents, such as carbon dioxide (CO 2 ), to reduce unwanted fish passage without disrupting human use. The purpose of this study was to evaluate the behavioral responses of common carp ( Cyprinus carpio ) to non-physical deterrents within a navigation structure on the Fox River, Wisconsin. Acoustic telemetry combined with hidden Markov models (HMMs) was used to analyze variation in carp responses to treatments. Outcomes may inform CO 2 effectiveness at preventing invasive carp movement through movement pinch-points.

Wisconsin

The importance of scaling for detecting community patterns: success and failure in assemblages of introduced species

Community saturation can help to explain why biological invasions fail. However, previous research has documented inconsistent relationships between failed invasions ( i.e ., an invasive species colonizes but goes extinct) and the number of species present in the invaded community. We use data from bird communities of the Hawaiian island of Oahu, which supports a community of 38 successfully established introduced birds and where 37 species were introduced but went extinct (failed invasions). We develop a modified approach to evaluate the effects of community saturation on invasion failure. Our method accounts (1) for the number of species present (NSP) when the species goes extinct rather than during its introduction; and (2) scaling patterns in bird body mass distributions that accounts for the hierarchical organization of ecosystems and the fact that interaction strength amongst species varies with scale. We found that when using NSP at the time of extinction, NSP was higher for failed introductions as compared to successful introductions, supporting the idea that increasing species richness and putative community saturation mediate invasion resistance. Accounting for scale-specific patterns in body size distributions further improved the relationship between NSP and introduction failure. Results show that a better understanding of invasion outcomes can be obtained when scale-specific community structure is accounted for in the analysis.

Hawaii

Trends in nonindigenous aquatic species richness in the United States reveal shifting spatial and temporal patterns of species introductions

Understanding the spatial and temporal dynamics underlying the introduction and spread of nonindigenous aquatic species (NAS) can provide important insights into the historical drivers of biological invasions and aid in forecasting future patterns of nonindigenous species arrival and spread. Increasingly, public databases of species observation records are being used to quantify changes in NAS distributions across space and time, and are becoming an important resource for researchers, managers, and policy-makers. Here we use publicly available data to describe trends in NAS introduction and spread across the conterminous United States over more than two centuries of observation records. Available data on first records of NAS reveal significant shifts in dominance of particular introduction patterns over time, both in terms of recipient regions and likely sources. These spatiotemporal trends at the continental scale may be subject to biases associated with regional variation in sampling effort, reporting, and data curation. We therefore also examined two additional metrics, the number of individual records and the spatial coverage of those records, which are likely to be more closely associated with sampling effort. Our results suggest that broad-scale patterns may mask considerable variation across regions, time periods, and even entities contributing to NAS sampling. In some cases, observed temporal shifts in species discovery may be influenced by dramatic fluctuations in the number and spatial extent of individual observations, reflecting the possibility that shifts in sampling effort may obscure underlying rates of NAS introduction.

Aquatic Invasions

Modeling habitat distribution from organism occurrences and environmental data: Case study using anemonefishes and their sea anemone hosts

We demonstrate the KGSMapper (Kansas Geological Survey Mapper), a straightforward, web-based biogeographic tool that uses environmental conditions of places where members of a taxon are known to occur to find other places containing suitable habitat for them. Using occurrence data for anemonefishes or their host sea anemones, and data for environmental parameters, we generated maps of suitable habitat for the organisms. The fact that the fishes are obligate symbionts of the anemones allowed us to validate the KGSMapper output: we were able to compare the inferred occurrence of the organism to that of the actual occurrence of its symbiont. Characterizing suitable habitat for these organisms in the Indo-West Pacific, the region where they naturally occur, can be used to guide conservation efforts, field work, etc.; defining suitable habitat for them in the Atlantic and eastern Pacific is relevant to identifying areas vulnerable to biological invasions. We advocate distinguishing between these 2 sorts of model output, terming the former maps of realized habitat and the latter maps of potential habitat. Creation of a niche model requires adding biotic data to the environmental data used for habitat maps: we included data on fish occurrences to infer anemone distribution and vice versa. Altering the selection of environmental variables allowed us to investigate which variables may exert the most influence on organism distribution. Adding variables does not necessarily improve precision of the model output. KGSMapper output distinguishes areas that fall within 1 standard deviation (SD) of the mean environmental variable values for places where members of the taxon occur, within 2 SD, and within the entire range of values; eliminating outliers or data known to be imprecise or inaccurate improved output precision mainly in the 2 SD range and beyond. Thus, KGSMapper is robust in the face of questionable data, offering the user a way to recognize and clean such data. It also functions well with sparse datasets. These features make it useful for biogeographic meta-analyses with the diverse, distributed datasets that are typical for marine organisms lacking direct commercial value. ?? Inter-Research 2006.

Marine Ecology Progress Series

Potential utility of environmental DNA for early detection of Eurasian watermilfoil ( Myriophyllum spicatum )

Considering the harmful and irreversible consequences of many biological invasions, early detection of an invasive species is an important step toward protecting ecosystems (Sepulveda et al. 2012). Early detection increases the probability that suppression or eradication efforts will be successful because invasive populations are small and localized (Vander Zanden et al. 2010). However, most invasive species are not detected early because current tools have low detection probabilities when target species are rare and the sampling effort required to achieve acceptable detection capabilities with current tools is seldom tractable (Jerde et al. 2011). As a result, many invasive species go undetected until they are abundant and suppression efforts become costly. Novel DNA-based surveillance tools have recently revolutionized early detection abilities using environmental DNA (eDNA) present in the water (Darling and Mahon 2011, Bohmann et al. 2014). In brief, eDNA monitoring enables the identification of organisms from DNA present and collected in water samples. Aquatic and semiaquatic organisms release DNA contained in sloughed, damaged, or partially decomposed tissue and waste products into the water and molecular techniques allow this eDNA in the water column to be identified from simple and easy-tocollect water samples (Darling and Mahon 2011). Despite limited understanding of the production, persistence, and spread of DNA in water (Barnes et al. 2014), eDNA monitoring has been applied not only to invasive species (Jerde et al. 2011), but also to species that are rare, endangered, or highly elusive (Spear et al. 2014). However, most eDNA research and monitoring has focused on detection of invertebrates and vertebrates and less attentionhas been given to developing eDNA techniques for detecting aquatic invasive plants. Eurasian watermilfoil (EWM; Myriophyllum spicatum L.) is an invasive species for which improved early detection would be particularly helpful. Advanced EWM invasions have negative impacts on native biodiversity, recreational boating, fishing, and other types of aquatic tourism (e.g., Eiswerth et al. 2000). On a broader scale, EWM can also be harmful to man-made aquatic infrastructure, such as hydroelectric dams. If an EWM invasion can be detected in an early stage where eradication is still a possibility, many of these negative consequences can be limited or prevented altogether (e.g., Madsen et al. 2002). The purpose of this research was to develop and validate a traditional polymerase chain reaction (PCR) assay for the detection of pure and hybridized EWM DNA using both laboratory and field experiments. We performed a pilot experiment in outdoor tanks to determine the basic functionality and sensitivity of the assay. Following this initial test, we collected field samples from Michigan and Montana lakes with and without known EWM populations. Taken together, our findings suggest that eDNA techniques have potential to be a useful strategy for the early detection of EWM.

Journal of Aquatic Plant Management

Moving from eco-forecasts to eco-projections

Ecological models can provide estimates of future conditions that are useful for decision-making, including long-term planning and resource prioritization. However, these models often rely on assumptions about ecological relationships and trajectories, forcings (e.g., biophysical conditions), and management approaches that may not be explicitly considered. To make assumptions more transparent, disciplines such as economics, demographics, climatology, and national intelligence make a fairly clear and consistent distinction between “forecasts” and “projections”. Forecasts are typically more near-term and rely on extending existing relationships and trends to estimate the most likely future conditions; whereas projections evaluate conditions under multiple scenarios that are based on an array of assumptions, often going further out in time. Consistently referring to ecological models of future conditions as either “eco-forecasts” or “eco-projections” could help make modelling assumptions more transparent and thus more effectively focus their application across landscapes and through time. To the extent that ecological modelling is used to support management, policy, and programmatic decisions, practitioners can ask the following. If the modelling is an eco-forecast, is it worth considering different initial conditions, trajectories, or forcings based on alternative scenarios? If the modelling is an eco-projection, are the underlying assumptions and future scenarios explicit, and are decisions properly tempered with respect to those modelling specifications? We demonstrate these concepts and methods for conducting eco-projections through examples from invasion biology and climate adaptation.

Conference Paper

Restoring Pacific Lamprey in the Umpqua River Basin of Oregon: A workshop summary

The Umpqua River Basin in southwestern Oregon (Figure 1) is part of the lands inhabited by the Cow Creek Band of Umpqua Tribe of Indians and an area of active co-management authority. This Basin supports a unique fish fauna, including important populations of Pacific salmon ( Oncorhynchus spp.) and steelhead ( O. mykiss ), and other native fishes that are endemic to the region (Mims et al. 2018). Among these species, the Pacific Lamprey ( Entosphenus tridentatus ) is one of the most unique, representing an ancient lineage of jawless fishes that long predates the evolution of any other species of fish in the basin (Clemens et al. 2017, 2021a). The Pacific Lamprey also represents an important cultural and food resource that features prominently in the indigenous practices of the Cow Creek Band of Umpqua Tribe of Indians. This report provides a select summary of topics related to the conservation of Pacific Lamprey in the Umpqua River Basin. Many of the topics addressed herein were discussed in a workshop co-hosted by the Cow Creek Band of Umpqua Tribe of Indians and the U.S. Geological Survey in April of 2022. This workshop was focused on threats to Pacific Lamprey in fresh waters of the basin. The workshop highlighted science conducted over the past 15 years by local managers and researchers to understand and address these threats and provide new information relevant to the conservation and restoration of Pacific Lamprey. Attendees included staff from the Coquille Indian Tribe, Yakama Nation, Columbia River Inter-Tribal Fish Commission, Partnership for Umpqua Rivers, Rogue River Watershed Council, Curry Watersheds Council, PacifiCorp, Trout Unlimited, Jackson Soil and Water Conservation District, Oregon Department of Fish and Wildlife, Bureau of Land Management, U.S. Fish and Wildlife Service, and U.S. Forest Service. This report provides an overview of this workshop and recent science and provides an overview of potential future efforts that could inform restoration of Pacific Lamprey. This summary is organized into four sections that relate to the main topics of the workshop: 1) The physical habitat template (stream flow, instream wood, sediment, and water temperature), 2) movement barriers and reservoirs, 3) biological invasions, and 4) climate adaptation. These do not represent an exhaustive list of topics related to conservation of Pacific Lamprey in the Umpqua Basin.

Oregon

Simulation of post-hurricane impact on invasive species with biological control management

Understanding the effects of hurricanes and other large storms on ecological communities and the post-event recovery in these communities can guide management and ecosystem restoration. This is particularly important for communities impacted by invasive species, as the hurricane may affect control efforts. Here we consider the effect of a hurricane on tree communities in southern Florida that has been invaded by Melaleuca quinquevervia (melaleuca), an invasive Australian tree. Biological control agents were introduced starting in the 1990s and are reducing melaleuca in habitats where they are established. We used size-structured matrix modeling as a tool to project the continued possible additional effects of a hurricane on a pure stand of melaleuca that already had some level of biological control. The model results indicate that biological control could suppress or eliminate melaleuca within decades. A hurricane that does severe damage to the stand may accelerate the trend toward elimination of melaleuca with both strong and moderate biological control. However, if the biological control is weak, the stand is resilient to all but extremely severe hurricane damage. Although only a pure melaleuca stand was simulated in this study, other plants, such as natives, are likely to accelerate the decline of melaleuca due to competition. Our model provides a new tool to simulate post-hurricanes effect on invasive species and highlights the essential role that biological control has played on invasive species management.

Florida

Risk analysis for biological hazards: What we need to know about invasive species

Risk analysis for biological invasions is similar to other types of natural and human hazards. For example, risk analysis for chemical spills requires the evaluation of basic information on where a spill occurs; exposure level and toxicity of the chemical agent; knowledge of the physical processes involved in its rate and direction of spread; and potential impacts to the environment, economy, and human health relative to containment costs. Unlike typical chemical spills, biological invasions can have long lag times from introduction and establishment to successful invasion, they reproduce, and they can spread rapidly by physical and biological processes. We use a risk analysis framework to suggest a general strategy for risk analysis for invasive species and invaded habitats. It requires: (1) problem formation (scoping the problem, defining assessment endpoints); (2) analysis (information on species traits, matching species traits to suitable habitats, estimating exposure, surveys of current distribution and abundance); (3) risk characterization (understanding of data completeness, estimates of the “potential” distribution and abundance; estimates of the potential rate of spread; and probable risks, impacts, and costs); and (4) risk management (containment potential, costs, and opportunity costs; legal mandates and social considerations and information science and technology needs).

Risk Analysis

Combined effects of biological control of an invasive shrub and fluvial processes on riparian vegetation dynamics

Plant community responses to biocontrol of invasive plants are understudied, despite the strong influence of the composition of replacement vegetation on ecosystem functions and services. We studied the vegetation response to a folivore beetle ( Diorhabda genus, Coleoptera) that has been introduced along southwestern US river valleys to control the invasion of non-native shrubs in the genus Tamarix (Tamaricaceae). We collected detailed plant compositional and environmental data during four different surveys over 7 years (2010–2017), including two surveys prior to when substantial beetle-induced dieback occurred in summer 2012, along the lower Virgin River, Nevada. The study river was of special interest because it is one of only a few largely unregulated rivers in the region, and a large flood of 40-year return period occurred between the first and second surveys, allowing us to study the combined effects of fluvial processes, which typically drive riparian plant community assembly, and biocontrol. Vegetation trajectories differed as a function of the dominant geomorphological process. Tamarix cover declined an average of 75% and was replaced by the native shrub Pluchea sericea as the new dominant species in the floodplain, especially where sediment deposition predominated. Following deposition, and especially erosion, opportunistic native herbs, Tamarix seedlings, and noxious weeds colonized the understory layer but did not increase in cover over time. Stands of the native shrub Salix exigua , a desirable replacement species following Tamarix control, only increased slightly and remained subordinate in the floodplain. Overall, our results showed that, by successfully controlling the target non-native plant, a biocontrol agent can substantially modify the replacement plant communities in a riparian system, but that fluvial processes also strongly influence the resulting communities.

Arizona, Nevada, Utah