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Modeling, mapping, and measuring the risk of freshwater invasive species across Alaska

Freshwater ecosystems of the Alaskan Arctic and Subarctic provide resources that are culturally, ecologically, and economically invaluable. Presently, these regions are relatively free of the impacts from invasive species compared to southern latitudes. To date, there have been relatively few verified introductions of aquatic invasive species (AIS) to freshwater ecosystems in Alaska. The expanding list and distribution of AIS has led to significant negative ecological and economic impacts (e.g., waterweed Elodea nuttalli ; E. canadensis and northern pike Esox Lucius introduced outside its native range in Alaska). Escalating human activity across Alaskan lands and waters, coupled with rapidly shifting environmental conditions, increases the potential for new species introductions and subsequent establishment. Creating a proactive framework for well-informed decision-making and action can improve the effectiveness of prevention efforts and bolster decision support tools that help resource managers direct limited resources. Prioritizing AIS that may be introduced and become established, as well as the locations at highest risk of invasion, is foundational to building a proactive invasive species management framework in Alaska. This project sought to identify and prioritize AIS known to be invasive in the contiguous United States, evaluate current and future habitat suitability for AIS in Alaska, and assess potential for AIS to be transported to habitats across Alaska, utilizing similar assessment methods as implemented for Bering Sea marine invasive species and non-native plants in Alaska. To accomplish this goal, the objectives of the project were to: 1) develop a formal ranked list of potential AIS to freshwater systems of Alaska; 2) assess the level of establishment risk for potential AIS by developing habitat suitability models for waterbodies across Alaska; and 3), identify potential pathways and specific vectors for high-risk AIS to invade Alaska and develop a framework for how vector analysis will be completed to understand transport risk. Overall, our goal is horizon scanning which is defined by Roy et al. (2019) as “a systematic examination of potential threats and opportunities, within a given context, and likely future developments, which are at the margin of current thinking and planning.” The scans include pathway analyses and risk screening of species present at pathway origin points, with a focus on identifying species at high risk of being introduced, becoming established, spreading, and causing harm. We refined a list of 28 AIS from a list of hundreds based on characterizations of species’ invasiveness and species’ proximity to Alaska (USGS 2020; GBIF 2022). Next, we evaluated the relative invasiveness of individual species to create an initial AIS ranking. We sought to characterize habitat suitability of AIS by selecting variables that were continental in scale, covering North America to include Alaska as well as the lower 48 states comparing natural discharge, sub-basin average terrain slope (degrees), average silt fraction, average organic carbon, lithological class, and human footprint in sub-basin in 2009. We estimated AIS habitat suitability across the entire state of Alaska using the physiological tolerances of the AIS (Appendix 2). We also evaluated pathways and vectors for the introduction of AIS (Appendix 2). Many pathways and vectors considered did not meet the criteria for Alaska or freshwater systems. Of the 28 ranked species that we categorized as very high, high, and moderate levels of invasiveness; all three risk groups included fish and mollusks (Appendix 2). One commonality of the very high-invasiveness-ranked species was the availability of Ecological Risk Screening Summary documents (USFWS, 2022) produced by U.S. Fish and Wildlife Service (USFWS), except for the goldfish ( Carassius auratus ) and the New Zealand mudsnail ( Potamopyrgus antipodarum ). The Ecological Risk Screening Summary is now available for New Zealand mudsnails. In general, fish species often ranked very high or high in invasiveness and included sportfish and aquarium fish, suggesting the importance of pathways such as aquarium trade, fishing industry, intentional (but illegal) introductions of sportfishes and aquarium fishes for establishment. The technique we used for habitat suitability models necessitated aquatic environmental datasets that were continental in scale, which was often interpolated from very coarse resolution source data layers, particularly in Alaska. Better spatial data representing aquatic environments would likely improve this approach. While the lack of introductions in Alaska and nearby provinces and states is encouraging, the lack of occurrence data for the focal species also created complications for habitat suitability modeling. Despite the challenges, the habitat suitability models indicated limited suitability for warmwater species while some species, such as Brook trout (S alvelinus fontinalis ), have high habitat suitability across Alaska no matter what threshold approach is taken. Some environmental predictors were more important than others. Specifically, the most important predictor variable, ‘frost free days,’ was critical for 15 out of 28 species as expected due to harsh winter conditions in Arctic and Subarctic regions. The second most important predictor was ‘subbasin land surface runoff’, a variable that indicates the amount of discharge and runoff, while the third most important predictor was ‘snow cover’ another indication of winter conditions. Overall, the ability to understand the effect of future climate scenarios on the establishment of AIS was challenging. A detailed dataset of freshwater temperatures and water chemistry (e.g., pH, calcium) would greatly improve the ability to predict invasiveness of freshwater species to Alaska’s ecosystems on a regional basis. Future studies may benefit from a more focused geographic scope examining a group of subbasins or a regional basin rather than the entire state. These drainages could be selected based upon the mostly likely locations of introduction pathways. The two most prevalent pathway risks for AIS are in-state transfer and stowaways/contaminants. Although there are examples of introductions from other pathways, the risk is somewhat mitigated by Alaska’s climate and regulations. However, variable application of protocols for inspection and cleaning of fishing gear, watercraft, and other similar items while traveling into Alaska as well as transferring from waterbody to waterbody within the state creates a substantial risk in introducing invasive species. We plot cumulative invasive vulnerability for all subbasins and for the top 10% of subbasins (Appendix 3).

Alaska↗

Potential spread of cheatgrass (Bromus tectorum) by feral horses (Equus ferus caballus) in Western Colorado

The invasive grass cheatgrass ( Bromus tectorum L.) presents major challenges for land management and habitat conservation in the western United States. Feral horses (Equus ferus caballus) have become overabundant in some areas of the West and can impact fragile semiarid ecosystems. Amid ongoing efforts to control cheatgrass in the Great Basin, we conducted a study to determine if feral horses contribute to the spread of cheatgrass through distribution via their feces. We collected feral horse fecal samples from Little Book Cliffs Herd Management Area in western Colorado in 2014. Fecal samples were dried, and 20 from each of 3 collection sessions were cultivated to examine germination success. Six species germinated from 18 samples (30%; mostly one plant per sample where germination occurred), including cheatgrass from 8% of samples. In a separate study we examined the diet of this same horse population using fecal plant DNA barcoding. Plant species that germinated were rare in the diet and germinated from fewer samples than expected relative to their detection in the diet. Our results suggest that feral horses could be contributing to cheatgrass propagation. Native ungulates and domestic cattle also have this potential. Although management of all large ungulates is necessary to mitigate cheatgrass spread, control of feral horse numbers is particularly necessary.

Colorado↗

Soil nutrients influence spatial distributions of tropical trees species

The importance of niche vs. neutral assembly mechanisms in structuring tropical tree communities remains an important unsettled question in community ecology [Bell G (2005) Ecology 86:1757-1770]. There is ample evidence that species distributions are determined by soils and habitat factors at landscape (<104 km2) and regional scales. At local scales (<1 km2), however, habitat factors and species distributions show comparable spatial aggregation, making it difficult to disentangle the importance of niche and dispersal processes. In this article, we test soil resource-based niche assembly at a local scale, using species and soil nutrient distributions obtained at high spatial resolution in three diverse neotropical forest plots in Colombia (La Planada), Ecuador (Yasuni), and Panama (Barro Colorado Island). Using spatial distribution maps of >0.5 million individual trees of 1,400 species and 10 essential plant nutrients, we used Monte Carlo simulations of species distributions to test plant-soil associations against null expectations based on dispersal assembly. We found that the spatial distributions of 36-51% of tree species at these sites show strong associations to soil nutrient distributions. Neutral dispersal assembly cannot account for these plant-soil associations or the observed niche breadths of these species. These results indicate that belowground resource availability plays an important role in the assembly of tropical tree communities at local scales and provide the basis for future investigations on the mechanisms of resource competition among tropical tree species. ?? 2007 by The National Academy of Sciences of the USA.

Proceedings of the National Academy of Sciences of↗

A test of the Niche Variation Hypothesis in a ruminant herbivore

Despite the shared prediction that the width of a population's dietary niche expands as food becomes limiting, the Niche Variation Hypothesis (NVH) and Optimal Foraging Theory (OFT) offer contrasting views about how individuals alter diet selection when food is limited. Classical OFT predicts that dietary preferences do not change as food becomes limiting, so individuals expand their diets as they compensate for a lack of preferred foods. In contrast, the NVH predicts that among-individual variation in cognition, physiology or morphology create functional trade-offs in foraging efficiency, thereby causing individuals to specialize on different subsets of food as food becomes limiting. To evaluate (a) the predictions of the NVH and OFT and (b) evidence for physiological and cognitive-based functional trade-offs, we used DNA microsatellites and metabarcoding to quantify the diet, microbiome and genetic relatedness (a proxy for social learning) of 218 moose Alces alces across six populations that varied in their degree of food limitation. Consistent with both the NVH and OFT, dietary niche breadth increased with food limitation. Increased diet breadth of individuals—rather than increased diet specialization—was strongly correlated with both food limitation and dietary niche breadth of populations, indicating that moose foraged in accordance with OFT. Diets were not constrained by inheritance of the microbiome or inheritance of diet selection, offering support for the little-tested hypothesis that functional trade-offs in food use (or lack thereof) determine whether populations adhere to the predictions of the NVH or OFT. Our results indicate that both the absence of strong functional trade-offs and the digestive physiology of ruminants provide contexts under which populations should forage in accordance with OFT rather than the NVH. Also, because dietary niche width increased with increased food limitation, OFT and the NVH provide theoretical support for the notion that plant–herbivore interaction networks are plastic rather than static, which has important implications for understanding interspecific niche partitioning. Lastly, because population-level dietary niche breadth and calf recruitment are correlated, and because calf recruitment can be a proxy for food limitation, our work demonstrates how diet data can be employed to understand a populations' proximity to carrying capacity.

Colorado, Utah, Wyoming↗

Adaptive management assists reintroduction as higher tides threaten an endangered salt marsh plant

In theory, extirpated plant species can be reintroduced and managed to restore sustainable populations. However, few reintroduced plants are known to persist for more than a few years. Our adaptive‐management case study illustrates how we restored the endangered hemiparasitic annual plant, Chloropyron maritimum subsp. maritimum (salt marsh bird's beak), to Sweetwater Marsh, San Diego Bay National Wildlife Refuge, California, United States, and used monitoring and experimentation to identify the factors limiting the reintroduced population. After extirpation in 1988, reintroduction starting that year led to a resilient, genetically diverse population in 2016 (a “boom” of approximately 14,000) that rebounded from a “bust” (62 in 2014). Multiple regressions attributed 82% of the variation in population counts to tidal amplitude, rainfall, and temperature. Populations of salt marsh bird's beak crashed when the diurnal tide range peaked during the 18.6‐year lunar nodal cycle (a rarely considered factor that periodically added approximately 12 cm to tidal ranges). We explain booms as follows: During smaller tidal amplitudes , above‐average rainfall could desalinize upper intertidal soils and stimulate salt marsh bird's beak germination. Then, moderate temperature in May favors growth to reproduction in June. In addition, salt marsh bird's beak needs a short and open canopy of native perennial plants, with roots to parasitize (not non‐native annual grass pseudohosts) and nearby upland soil for a preferred pollinator, ground‐burrowing bees. Although our reintroduced salt marsh bird's beak population is an exceptional case of persistence, this rare species‐specific environmental and biological requirement makes it vulnerable to rising sea levels and global warming.

California↗

Habitat suitability and ecological associations of two non-native ungulate species on the Hawaiian island of Lanai

The ability to effectively manage game species for specific conservation objectives is often limited by the scientific understanding of their distribution and abundance. This is especially true in Hawai‘i where introduced game mammals are poorly studied and have low value relative to native species in other states. We modeled the habitat suitability and ecological associations of European mouflon sheep (“mouflon”; Ovis musimon ) and axis deer ( Axis axis ) on the island of Lāna‘i using intensive aerial survey and environmental data that included climate, vegetation, and topographic variables. We conducted diagnostic tests on a suite of primarily categorical predictors and determined most were highly correlated. We therefore developed a suite of other spatial predictor layers with continuous variables. We tested several modeling approaches but settled on generalized linear models (GLM) and random GLMs because they could account for group size of animals and were based on curvilinear responses of each species to environmental variability. Both mammal species were habitat generalists showing little affinity to particular plant species or communities. Continuous predictors associated with plant productivity such as mean annual precipitation, normalized difference vegetation index (NDVI), and cloud cover were important explanatory factors in a GLM of axis deer and a random GLM of mouflon habitat suitability. The presence of axis deer was also an important explanatory predictor for mouflon distribution, but deer were not influenced by mouflon distribution, indicating asymmetrical competition. Consequently, mouflon were restricted to lower elevation arid and very dry slopes, whereas axis deer were more broadly distributed throughout other upland environments of the island, but avoided steep terrain. Findings indicate that removal of a substantial portion of the more abundant axis deer population may lead to an increase in abundance and distribution of mouflon without containment. Resulting spatial models of game mammal habitat suitability will be employed to inform land use prioritization analyses and to help resolve long-standing conflicts between native species conservation and sustained-yield hunting.

Hawai'i↗

Food web heterogeneity and succession in created saltmarshes

1. Ecological restoration must achieve functional as well as structural recovery. Functional metrics for reestablishment of trophic interactions can be used to complement traditional monitoring of structural attributes. In addition, topographic effects on food web structure provide added information within a restoration context; often, created sites may require spatial heterogeneity to effectively match structure and function of natural habitats. 2. We addressed both of these issues in our study of successional development of benthic food web structure, with focus on bottom&ndash;up driven changes in macroinvertebrate consumer assemblages in the salt marshes of the Venice Lagoon, Italy. We combined quantified estimates of the changing community composition with stable isotope data (13C:12C and 15N:14N) to compare the general trophic structure between created (2&ndash;14 years) marshes and reference sites and along topographic elevation gradients within salt marshes. 3. Macrofaunal invertebrate consumers exhibited local, habitat-specific trophic patterns. Stable isotope-based trophic structure changed with increasing marsh age, in particular with regards to mid-elevation (Salicornia) habitats. In young marshes, the mid-elevation consumer signatures resembled those of unvegetated ponds. The mid elevation of older and natural marshes had a more distinct Salicornia-zone food web, occasionally resembling that of the highest (Sarcocornia-dominated) elevation. In summary, this indicates that primary producers and availability of vascular plant detritus structure consumer trophic interactions and the flow of carbon. 4. Functionally different consumers, subsurface-feeding detritivores (Oligochaeta) and surface grazers (Hydrobia sp.), showed distinct but converging trajectories of isotopic change over time, indicating that successional development may be asymmetric between &lsquo;brown&rsquo; (detrital) guilds and &lsquo;green&rsquo; (grazing) guilds in the food web. 5. Synthesis and applications. Created marsh food webs converged into a natural state over about a decade, with successional shifts seen in both consumer community composition and stable isotope space. Strong spatial effects were noted, highlighting the utility of stable isotopes to evaluate functional equivalence in spatially heterogeneous systems. Understanding the recovery of functional properties such as food web support, and their inherent spatial variability, is key to planning and managing successful habitat restoration.

Adriatic Sea, Venice Lagoon↗

Widespread use of the nitrification inhibitor nitrapyrin: Assessing benefits and costs to agriculture, ecosystems, and environmental health

Agricultural production and associated applications of nitrogen (N) fertilizers have increased dramatically in the last century, and current projections to 2050 show that demands will continue to increase as the human population grows. Applied in both organic and inorganic fertilizer forms, N is an essential nutrient in crop productivity. Increased fertilizer applications, however, create the potential for more N loss before plant uptake. One strategy for minimizing N loss is the use of enhanced efficiency fertilizers, fortified with a nitrification inhibitor, such as nitrapyrin. In soils and water, nitrapyrin inhibits the activity of ammonia monooxygenase, a microbial enzyme that catalyzes the first step of nitrification from ammonium to nitrite. Potential benefits of using nitrification inhibitors range from reduced nitrate leaching and nitrous oxide emissions to increased crop yield. The extent of these benefits, however, depends on environmental conditions and management practices. Thus, such benefits are not always realized. Additionally, nitrapyrin has been shown to transport off-field, and it is unknown what effects environmental nitrapyrin could have on nontarget organisms and the ecological nitrogen cycle. Here, we review the agronomic and environmental benefits and costs of nitrapyrin use and present a series of research questions and considerations to be addressed with future nitrification inhibitor research.

Environmental Science and Technology↗

Estimating vegetation biomass and cover across large plots in shrub and grass dominated drylands using terrestrial lidar and machine learning

Terrestrial laser scanning (TLS) has been shown to enable an efficient, precise, and non-destructive inventory of vegetation structure at ranges up to hundreds of meters. We developed a method that leverages TLS collections with machine learning techniques to model and map canopy cover and biomass of several classes of short-stature vegetation across large plots. We collected high-definition TLS scans of 26 1-ha plots in desert grasslands and big sagebrush shrublands in southwest Idaho, USA. We used the Random Forests machine learning algorithm to develop decision tree models predicting the biomass and canopy cover of several vegetation classes from statistical descriptors of the aboveground heights of TLS points. Manual measurements of vegetation characteristics collected within each plot served as training and validation data. Models based on five or fewer TLS descriptors of vegetation heights were developed to predict the canopy cover fraction of shrubs (R 2 = 0.77, RMSE = 7%), annual grasses (R 2 = 0.70, RMSE = 21%), perennial grasses (R 2 = 0.36, RMSE = 12%), forbs (R 2 = 0.52, RMSE = 6%), bare earth or litter (R 2 = 0.49, RMSE = 19%), and the biomass of shrubs (R 2 = 0.71, RMSE = 175 g) and herbaceous vegetation (R 2 = 0.61, RMSE = 99 g) (all values reported are out-of-bag). Our models explained much of the variability between predictions and manual measurements, and yet we expect that future applications could produce even better results by reducing some of the methodological sources of error that we encountered. Our work demonstrates how TLS can be used efficiently to extend manual measurement of vegetation characteristics from small to large plots in grasslands and shrublands, with potential application to other similarly structured ecosystems. Our method shows that vegetation structural characteristics can be modeled without classifying and delineating individual plants, a challenging and time-consuming step common in previous methods applying TLS to vegetation inventory. Improving application of TLS to studies of shrub-steppe ecosystems will serve immediate management needs by enhancing vegetation inventories, environmental modeling studies, and the ability to train broader datasets collected from air and space.

Idaho↗

Human infrastructure and invasive plant occurrence across rangelands of southwestern Wyoming, U.S.A.

Although human influence across rural landscapes is often discussed, interactions between the native, natural systems and human activities are challenging to measure explicitly. We assessed the distribution of introduced, invasive species as related to anthropogenic infrastructure and environmental conditions across southwestern Wyoming. to discern direct correlations as well as covariate influences between land use, land cover, and abundance of invasive plants, and assess the supposition that these features affect surrounding rangeland conditions. Our sample units were 1 000 m long and extended outward from target features, which included roads, oil and gas well pads, pipelines, power lines, and featureless background sites. Sample sites were distributed across the region using a stratified, random design with a frame that represented features and land-use intensity. In addition to land-use gradients, we captured a representative, but limited, range of variability in climate, soils, geology, topography, and dominant vegetation. Several of these variables proved significant, in conjunction with distance from anthropogenic features, in regression models of invasive plant abundance. We used general linear models to demonstrate and compare associations between invasive plant frequency and Euclidian distance from features, natural logarithm transformed distances (log-linear), and environmental variables which were presented as potential covariates. We expected a steep curvilinear (log or exponential) decline trending towards an asymptote along the axis representing high abundance near features with rapid decrease beyond approximately 50–100 m. Some of the associations we document exhibit this pattern, but we also found some invasive plant distributions that extended beyond our expectations, suggesting a broader distribution than anticipated. Our results provide details that can inform local efforts for management and control of invasive species, and they provide evidence of the different associations between natural patterns and human land use exhibited by nonnative species in this rural setting, such as the indirect effects of humans beyond impact areas.

Wyoming↗

Climate correlates of 20 years of trophic changes in a high-elevation riparian system

The consequences of climate change for ecosystem structure and function remain largely unknown. Here, I examine the ability of climate variation to explain long-term changes in bird and plant populations, as well as trophic interactions in a high-elevation riparian system in central Arizona, USA, based on 20 years of study. Abundances of dominant deciduous trees have declined dramatically over the 20 years, correlated with a decline in overwinter snowfall. Snowfall can affect overwinter presence of elk, whose browsing can significantly impact deciduous tree abundance. Thus, climate may affect the plant community indirectly through effects on herbivores, but may also act directly by influencing water availability for plants. Seven species of birds were found to initiate earlier breeding associated with an increase in spring temperature across years. The advance in breeding time did not affect starvation of young or clutch size. Earlier breeding also did not increase the length of the breeding season for single-brooded species, but did for multi-brooded species. Yet, none of these phenology-related changes was associated with bird population trends. Climate had much larger consequences for these seven bird species by affecting trophic levels below (plants) and above (predators) the birds. In particular, the climate-related declines in deciduous vegetation led to decreased abundance of preferred bird habitat and increased nest predation rates. In addition, summer precipitation declined over time, and drier summers also were further associated with greater nest predation in all species. The net result was local extinction and severe population declines in some previously common bird species, whereas one species increased strongly in abundance, and two species did not show clear population changes. These data indicate that climate can alter ecosystem structure and function through complex pathways that include direct and indirect effects on abundances and interactions of multiple trophic components. ?? 2007 by the Ecological Society of America.

Ecology↗

Potential cheatgrass abundance within lightly invaded areas of the Great Basin

Context Anticipating where an invasive species could become abundant can help guide prevention and control efforts aimed at reducing invasion impacts. Information on potential abundance can be combined with information on the current status of an invasion to guide management towards currently uninvaded locations where the threat of invasion is high. Objectives We aimed to support management by developing predictive maps of potential cover for cheatgrass ( Bromus tectorum ), a problematic invader that can transform plant communities. We integrated our predictions of potential abundance with mapped estimates of current cover to quantify invasion potential within lightly invaded areas. Methods We used quantile regression to model cheatgrass abundance as a function of climate, weather, and disturbance, treating outputs as low to high invasion scenarios. We developed a species-specific set of covariates and validated model performance using spatially and temporally independent data. Results Potential cheatgrass abundance was higher in areas that had burned, at low elevations, and when fall germination conditions were more favorable. Our results highlight the extensive areas across the Great Basin where cheatgrass abundance could increase to levels that can alter fire behavior and cause other ecological impacts. Conclusions We predict potential cheatgrass abundance to quantify relative invasion risk. Our model results provide high and low scenarios of cheatgrass abundance to guide resource allocation and planning efforts across shrubland ecosystems of the Great Basin that remain relatively uninvaded. Combining information on an invasive species’ current and potential abundance can yield spatial predictions to guide resource allocation and management action.

California, Idaho, Nevada, Oregon, Utah↗

Primary production responses to extreme changes in North American Monsoon precipitation vary by elevation and plant functional composition through time

Primary production in dryland ecosystems is limited by water availability and projected to be strongly affected by future shifts in seasonal precipitation. Warm-season precipitation derived from the North American Monsoon contributes 40% of annual precipitation to dryland ecosystems in the southwestern U.S. and is projected to become more variable. However, there is large uncertainty on whether this variability will be expressed as either extreme wet or dry years and how primary production of different plant functional types will respond across widespread elevation gradients in this region. We experimentally imposed extreme drought and water addition treatments from 2016 – 2020, during which ambient warm-season precipitation declined to reach historic lows, to understand production sensitivity of dominant plant functional types along a 1,000 m elevation gradient. We found that the production responses of plant functional types to monsoon precipitation extremes were dependent on the number of treatment years that occurred across sites along the elevation gradient. C 4 perennial grasses were most responsive to precipitation manipulation treatments, followed by C 3 perennial grasses and annuals, while perennial forbs and shrubs had weak or no responses. C 4 perennial grass reductions due to extreme drought were generally stronger or occurred earlier at low elevation sites, while multi-year extreme drought extended negative effects to C 3 perennial grasses at high elevation, and all sites showed delayed responses to multi-year water addition. We found that the sensitivity of C 3 perennial grass production differed for extreme drought and water addition compared to ambient precipitation at one site, but other sites and plant functional types had similar sensitivities to the different treatment types. Synthesis . The upward advance of primary production responsiveness from single- to multi-year extreme changes in warm-season precipitation suggests more immediate shifts in functional composition and carbon cycling at low elevation, while high elevation ecosystems may become less resistant as the effects of extreme precipitation compound through time.

Arizona↗

Persistence of native and exotic plants 10 years after prairie reconstruction

Prairie reconstructions are a critical component of preservation of the imperiled tallgrass prairie ecosystem in the Midwestern United States. Sustainability of this endeavor depends on establishment of persistent cover of planted native species and resistance to noxious weeds. The goal of this study was to understand the influence of early reconstruction practices on long-term outcomes. Twelve replicates of three planting methods (dormant-season broadcast, growing-season broadcast, and growing-season drill) and three seed mix richness levels (10, 20, or 34 species), fully crossed in a completely randomized design were planted in 2005 on nine former agricultural fields located in Iowa and Minnesota. Cover by species was estimated in 2005–2007, 2010, and 2015. In 2015, cover of planted species, native nonplanted species, and exotic species were similar to those recorded in 2010. Cover of the noxious weed Cirsium arvense had also declined by an average of 49% without herbicide from a peak in 2007 to low stable levels from 2010 to 2015. Richness of planted forbs, on the other hand, were still increasing in high-richness broadcast treatments (e.g. 17–59% increase 2010–1015 in Minnesota). Two results in 2015 are reasons for concern: cover of planted species is only slightly over 50% in both Minnesota and Iowa, though with forbs still increasing, this may improve; and the cool-season exotic grasses Poa pratensis and Bromus inermis are increasing at both Minnesota and Iowa sites. Control of these invasive grasses will be necessary, but care will be needed to avoid negative impacts of control methods on natives.

Restoration Ecology↗

Response of Everglades tree islands to environmental change

Tree islands are centers of biodiversity within the Florida Everglades, USA, but the factors controlling their distribution, formation, and development are poorly understood. We use pollen assemblages from tree islands throughout the greater Everglades ecosystem to reconstruct the timing of tree island formation, patterns of development, and response to specific climatic and environmental stressors. These data indicate that fixed (teardrop-shaped) and strand tree islands developed well before substantial human alteration of the system, with initial tree island vegetation in place between 3500 and 500 calibrated years before present (cal yr BP), depending on the location in the Everglades wetland. Tree island development appears to have been triggered by regional- to global-scale climatic events at 2800 cal yr BP, 1600–1500 cal yr BP, 1200–1000 cal yr BP (early Medieval Warm Period), and 500–200 cal yr BP (Little Ice Age). These periods correspond to drought intervals documented in Central and South America and periods of southward displacement of the Intertropical Convergence Zone. The records indicate a coherence of climate patterns in both subtropical North America and the Northern Hemisphere Neotropics. Water management practices of the 20th century altered plant communities and size of tree islands throughout the Everglades. Responses range from loss of tree islands due to artificially long hydroperiods and deep water to expansion of tree islands after flow reductions. These data provide evidence for the rapidity of tree island response to specific hydrologic change and facilitate prediction of the response to future changes associated with Everglades restoration plans.

Florida↗

Bay Miwok evening primrose: A new subspecies of Oenothera deltoides (Onagraceae) endemic to California

California contains exceptional biodiversity in geography and plant life, including numerous endemic species, some of which are cryptic. The Oenothera deltoides Torr. & Frém. species complex represents a prime example of cryptic diversity. Here, we recognize a new subspecies of Oenothera deltoides , O. deltoides subsp. julpunensis S.F.Jones, subsp. nov., that is a local endemic of windblown sand deposits on the eastern Antioch Dunes sand sheet in the San Francisco Bay-Delta region of California, USA. With the goal of providing clarity to managers of listed species and better understanding of California's diverse flora, we addressed the puzzle of O. deltoides in the region by combining range-wide field surveys with modern genomic tools. We describe the proposed subspecies, its ecology and distribution, and discuss its conservation. As a somewhat cryptic local endemic with small population size and disappearing habitat, the proposed subspecies would benefit from conservation and management to persist as a member of the California flora.

California↗

A review of natural and managed revegetation responses in two de-watered reservoirs after large dam removals on the Elwha River, Washington, USA

Large dam removals are increasing in frequency and the response of natural and managed revegetation is a critical consideration for managed restoration of dewatered reservoir landscapes post dam removal. The removal of two large dams on the Elwha River in 2011-2014 provides insight into reservoir revegetation. We review literature and datasets from 2012 through 2018, 1-6 years since reservoir dewatering, to compare pre-dam removal predictions on the Elwha to post-dam removal of natural revegetation, managed revegetation effects and invasive non-native vegetation response. Pre-dam removal hypotheses about natural revegetation did not predict species performance on reservoir sediments, seed rain patterns, or seed bank response. Sediment texture and landform affected multiple aspects of revegetation, including vegetation cover, species richness, woody stem densities and species composition. Reservoir drawdown timing influenced species composition and seedling densities. Predictions about managed revegetation effects were mixed. Planting trees and shrubs did not accelerate woody cover but did increase species richness. Seeding reduced non-native vegetation frequency and species richness, had no effect on vegetation cover on fine sediments, but increased vegetation cover on coarse sediments. Planting trees and shrubs during drawdown appeared to result in higher survival rates compared to plantings installed 1+ years post drawdown. Seeding Lupinus rivularis (riverbank lupine) on coarse sediments was successful and increased foliar nitrogen in planted conifers. Invasive non-native vegetation was correctly predicted to be more abundant in the Aldwell reservoir but did not preclude native species establishment in either reservoir, likely due to rapid establishment of native species and robust management that occurred before, during and after dam removal.

Washington↗

Feeding preferences of West Indian manatees in Florida, Belize, and Puerto Rico as indicated by stable isotope analysis

The endangered West Indian manatee Trichechus manatus has 2 recognized subspecies: the Florida T. m. latirostris and Antillean T. m. manatus manatee, both of which are found in freshwater, estuarine, and marine habitats. A better understanding of manatee feeding preferences and habitat use is essential to establish criteria on which conservation plans can be based. Skin from manatees in Florida, Belize, and Puerto Rico, as well as aquatic vegetation from their presumed diet, were analyzed for stable carbon and nitrogen isotope ratios. This is the first application of stable isotope analysis to Antillean manatees. Stable isotope ratios for aquatic vegetation differed by plant type (freshwater, estuarine, and marine), collection location, and in one instance, season. Carbon and nitrogen isotope ratios for manatee skin differed between collection location and in one instance, season, but did not differ between sex or age class. Signatures in the skin of manatees sampled in Belize and Puerto Rico indicated a diet composed primarily of seagrasses, whereas those of Florida manatees exhibited greater regional variation. Mixing model results indicated that manatees sampled from Crystal River and Homosassa Springs (Florida, USA) ate primarily freshwater vegetation, whereas manatees sampled from Big Bend Power Plant, Ten Thousand Islands, and Warm Mineral Springs (Florida) fed primarily on seagrasses. Possible diet–tissue discrimination values for 15 N were estimated to range from 1.0 to 1.5‰. Stable isotope analysis can be used to interpret manatee feeding behavior over a long period of time, specifically the use of freshwater vegetation versus seagrasses, and can aid in identifying critical habitats and improving conservation efforts.

Florida↗