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Twenty Years After the 1988 Yellowstone Fires: Lessons About Disturbance and Ecosystems

The 1988 Yellowstone fires were among the first in what has proven to be an upsurge in large severe fires in the western USA during the past 20 years. At the time of the fires, little was known about the impacts of such a large severe disturbance because scientists had had few previous opportunities to study such an event. Ecologists predicted short- and long-term effects of the 1988 fires on vegetation, biogeochemistry, primary productivity, wildlife, and aquatic ecosystems based on scientific understanding of the time. Twenty-plus years of subsequent study allow these early predictions to be evaluated. Most of the original predictions were at least partially supported, but some predictions were refuted, others nuanced, and a few postfire phenomena were entirely unexpected. Post-1988 Yellowstone studies catalyzed advances in ecology focused on the importance of spatial and temporal heterogeneity, contingent influences, and multiple interacting drivers. Post-1988 research in Yellowstone also has changed public perceptions of fire as an ecological process and attitudes towards fire management. Looking ahead to projected climate change and more frequent large fires, the well-documented ecological responses to the 1988 Yellowstone fires provide a foundation for detecting and evaluating potential changes in fire regimes of temperate mountainous regions. ?? 2011 Springer Science+Business Media, LLC.

Ecosystems

Practices of biological soil crust rehabilitation in China: Experiences and challenges

Biological soil crusts (biocrusts) are a central component of dryland ecosystems. However, they are highly vulnerable to disturbance and natural recovery may be slow. Therefore, finding ways to enhance the reestablishment of biocrusts after disturbance has been of great interest to researchers. This article provides a review of the laboratory cultivation and field inoculations of biocrust materials in China (mostly published in Chinese). Larger filamentous cyanobacteria (e.g. Microcoleus ) are relatively easy, although slow, to grow in culture compared to other biocrust components. Thus, most researchers have focused their efforts on the cyanobacteria and a few species of mosses that are also easily grown but at smaller scale. For all the studies, a small amount of biocrust material was collected and its biomass enhanced under controlled conditions. However, the enhancement was done using various methods and techniques in different regions. These materials were then applied to disturbed field sites, again with various methods. Results show that keeping the inoculated soil surface wet for some time period after inoculation was crucial for restoration success. Cyanobacterial establishment was improved by installing automatic sprinkling using micro‐irrigation techniques and/or physical structures that reduced sediment moving onto the inoculated area. Experimental applications in China showed that cyanobacteria can be successfully inoculated at a large scale (hundreds of ha). Moss inoculation, on the other hand, was only accomplished at a small scale (several m 2 ). To assess whether biocrust restoration can enhance the establishment of a self‐supporting ecosystem, further research is needed on how inoculation affects vegetation diversity and structure and ecological processes.

Restoration Ecology

Influence of offshore oil and gas structures on seascape ecological connectivity

Offshore platforms, subsea pipelines, wells and related fixed structures supporting the oil and gas (O&G) industry are prevalent in oceans across the globe, with many approaching the end of their operational life and requiring decommissioning. Although structures can possess high ecological diversity and productivity, information on how they interact with broader ecological processes remains unclear. Here, we review the current state of knowledge on the role of O&G infrastructure in maintaining, altering or enhancing ecological connectivity with natural marine habitats. There is a paucity of studies on the subject with only 33 papers specifically targeting connectivity and O&G structures, although other studies provide important related information. Evidence for O&G structures facilitating vertical and horizontal seascape connectivity exists for larvae and mobile adult invertebrates, fish and megafauna; including threatened and commercially important species. The degree to which these structures represent a beneficial or detrimental net impact remains unclear, is complex and ultimately needs more research to determine the extent to which natural connectivity networks are conserved, enhanced or disrupted. We discuss the potential impacts of different decommissioning approaches on seascape connectivity and identify, through expert elicitation, critical knowledge gaps that, if addressed, may further inform decision making for the life cycle of O&G infrastructure, with relevance for other industries (e.g. renewables). The most highly ranked critical knowledge gap was a need to understand how O&G structures modify and influence the movement patterns of mobile species and dispersal stages of sessile marine species. Understanding how different decommissioning options affect species survival and movement was also highly ranked, as was understanding the extent to which O&G structures contribute to extending species distributions by providing rest stops, foraging habitat, and stepping stones. These questions could be addressed with further dedicated studies of animal movement in relation to structures using telemetry, molecular techniques and movement models. Our review and these priority questions provide a roadmap for advancing research needed to support evidence-based decision making for decommissioning O&G infrastructure.

Global Change Biology

A quantitative framework for estimating risk of collision between marine mammals and boats

Speed regulations of watercraft in protected areas are designed to reduce lethal collisions with wildlife but can have economic consequences. We present a quantitative framework for investigating the risk of deadly collisions between boats and wildlife. We apply encounter rate theory to demonstrate how marine mammal-boat encounter rate can be used to predict the expected number of deaths associated with management scenarios. We illustrate our approach with management scenarios for two endangered species: the Florida manatee Trichechus manatus latirostris and the North Atlantic right whale Eubalaena glacialis . We used a Monte Carlo simulation approach to demonstrate the uncertainty that is associated with our estimate of relative mortality. We show that encounter rate increased with vessel speed but that the expected number of encounters varies depending on the boating activities considered. For instance, in a scenario involving manatees and boating activities such as water skiing, the expected number of encounters in a given area (in a fixed time interval) increased with vessel speed. In another scenario in which a vessel made a transit of fixed length the expected number of encounters decreases slightly with boat speed. In both cases the expected number of encounters increased with distanced travelled by the boat. For whales, we found a slight reduction (~0.1%) in the number of encounters under a scenario where speed is unregulated; this reduction, however, is negligible, and overall expected relative mortality was ~30% lower under the scenario with speed regulation. The probability of avoidance by the animal or vessel was set to 0 because of lack of data, but we explored the importance of this parameter on the model predictions. In fact, expected relative mortality under speed regulations decreases even further when the probability of avoidance is a decreasing function of vessel speed. By applying encounter rate theory to the case of boat collisions with marine mammals, we gained new insights about encounter processes between wildlife and watercraft. Our work emphasizes the importance of considering uncertainty when estimating wildlife mortality. Finally, our findings are relevant to other systems and ecological processes involving the encounter between moving agents.

Methods in Ecology and Evolution

Dynamic population models with temporal preferential sampling to infer phenology

To study population dynamics, ecologists and wildlife biologists typically use relative abundance data, which may be subject to temporal preferential sampling. Temporal preferential sampling occurs when the times at which observations are made and the latent process of interest are conditionally dependent. To account for preferential sampling, we specify a Bayesian hierarchical abundance model that considers the dependence between observation times and the ecological process of interest. The proposed model improves relative abundance estimates during periods of infrequent observation and accounts for temporal preferential sampling in discrete time. Additionally, our model facilitates posterior inference for population growth rates and mechanistic phenometrics. We apply our model to analyze both simulated data and mosquito count data collected by the National Ecological Observatory Network. In the second case study, we characterize the population growth rate and relative abundance of several mosquito species in the Aedes genus. Supplementary materials accompanying this paper appear on-line.

Journal of Agricultural, Biological, and Environme

Application of ecological criteria in selecting marine reserves and developing reserve networks

Marine reserves are being established worldwide in response to a growing recognition of the conservation crisis that is building in the oceans. However, designation of reserves has been largely opportunistic, or protective measures have been implemented (often overlapping and sometimes in conflict) by different entities seeking to achieve different ends. This has created confusion among both users and enforcers, and the proliferation of different measures provides a false sense of protection where little is offered. This paper sets out a procedure grounded in current understanding of ecological processes, that allows the evaluation and selection of reserve sites in order to develop functional, interconnected networks of fully protected reserves that will fulfill multiple objectives. By fully protected we mean permanently closed to fishing and other resource extraction. We provide a framework that unifies the central aims of conservation and fishery management, while also meeting other human needs such as the provision of ecosystem services (e.g., maintenance of coastal water quality, shoreline protection, and recreational opportunities). In our scheme, candidate sites for reserves are evaluated against 12 criteria focused toward sustaining the biological integrity and productivity of marine systems at both local and regional scales. While a limited number of sites will be indispensable in a network, many will be of similar value as reserves, allowing the design of numerous alternative, biologically adequate networks. Devising multiple network designs will help ensure that ecological functionality is preserved throughout the socioeconomic evaluation process. Too often, socioeconomic criteria have dominated the process of reserve selection, potentially undermining their efficacy. We argue that application of biological criteria must precede and inform socioeconomic evaluation, since maintenance of ecosystem functioning is essential for meeting all of the goals for reserves. It is critical that stakeholders are fully involved throughout this process. Application of the proposed criteria will lead to networks whose multifunctionality will help unite the objectives of different management entities, so accelerating progress toward improved stewardship of the oceans.

Ecological Applications

Increasing connectivity between metapopulation ecology and landscape ecology

Metapopulation ecology and landscape ecology aim to understand how spatial structure influences ecological processes, yet these disciplines address the problem using fundamentally different modeling approaches. Metapopulation models describe how the spatial distribution of patches affects colonization and extinction, but often do not account for the heterogeneity in the landscape between patches. Models in landscape ecology use detailed descriptions of landscape structure, but often without considering colonization and extinction dynamics. We present a novel spatially explicit modeling framework for narrowing the divide between these disciplines to advance understanding of the effects of landscape structure on metapopulation dynamics. Unlike previous efforts, this framework allows for statistical inference on landscape resistance to colonization using empirical data. We demonstrate the approach using 11 yr of data on a threatened amphibian in a desert ecosystem. Occupancy data for Lithobates chiricahuensis (Chiricahua leopard frog) were collected on the Buenos Aires National Wildlife Refuge (BANWR), Arizona, USA from 2007 to 2017 following a reintroduction in 2003. Results indicated that colonization dynamics were influenced by both patch characteristics and landscape structure. Landscape resistance increased with increasing elevation and distance to the nearest streambed. Colonization rate was also influenced by patch quality, with semi‐permanent and permanent ponds contributing substantially more to the colonization of neighboring ponds relative to intermittent ponds. Ponds that only hold water intermittently also had the highest extinction rate. Our modeling framework can be widely applied to understand metapopulation dynamics in complex landscapes, particularly in systems in which the environment between habitat patches influences the colonization process

Ecology

Mapping predicted ecological states at landscape scales using remote sensing data and machine learning

Dryland ecosystems, covering 45% of the Earth's land and supporting over one-third of the global population, face significant threats from land degradation and ecological state change. Managing these ecosystems is complex, and science-based frameworks like Ecological Site Descriptions and state-and-transition models are essential tools for guiding decisions to support ecological health while maintaining stakeholder values such as grazing, wildlife, and recreation. However, alignment of these frameworks with smaller scale soil survey maps limits their applicability to broader ecological processes. Here, we extend these frameworks to larger landscapes with a machine learning approach that integrates large-scale, high-resolution vegetation data with identified ecological states from a data-driven state-and-transition model developed for a landscape-scale Ecological Site Group. A “global” model, which used combined inputs from multiple remotely sensed datasets, outperformed individual dataset models based on evaluation with independent data. Ecological state maps generated through this approach broaden the utility of state-and-transition models across Ecological Site Groups, providing a more spatially robust tool for land management at watershed and larger landscape scales. These methods, and the associated ecological state maps, can help meet critical needs for improved land condition assessments that support development of resource management plans and help identify priority areas for restoration and conservation.

Arizona, Colorado, New Mexico, Utah, Wyoming

Mountain hydrology of the western United States

Climate change and climate variability, population growth, and land use change drive the need for new hydrologic knowledge and understanding. In the mountainous West and other similar areas worldwide, three pressing hydrologic needs stand out: first, to better understand the processes controlling the partitioning of energy and water fluxes within and out from these systems; second, to better understand feedbacks between hydrological fluxes and biogeochemical and ecological processes; and, third, to enhance our physical and empirical understanding with integrated measurement strategies and information systems. We envision an integrative approach to monitoring, modeling, and sensing the mountain environment that will improve understanding and prediction of hydrologic fluxes and processes. Here extensive monitoring of energy fluxes and hydrologic states are needed to supplement existing measurements, which are largely limited to streamflow and snow water equivalent. Ground‐based observing systems must be explicitly designed for integration with remotely sensed data and for scaling up to basins and whole ranges.

Water Resources Research

Establishing historical baselines of arthropod assemblages using rodent paleomiddens

Aim Arthropods are under-represented in paleoecological studies but are critical agents in ecological processes. Despite rigorous documentation of diverse arthropod assemblages from ancient rodent (or paleo) middens worldwide, their use for studying ancient arthropod diversity has stalled in recent decades. Here, we review published studies to identify how paleomidden arthropods can be leveraged to address significant questions in ecology and synthesise spatiotemporal trends in ancient arthropod diversity. Location Global with a focus on the Americas. Time Period From ~40,000 to 100 years before present. Major Taxa Studied Fossil arthropods preserved in rodent paleomiddens. Methods We identified four primary objectives in ecology that can be advanced using paleomidden arthropod fossils: the reconstruction of ancient biodiversity, the elucidation of mechanisms and processes driving arthropod range shifts, the incorporation of arthropods into ancient trophic webs, and the assessment of adaptive responses to changing environments. Results We identified 20 papers reporting arthropod remains, of which 14 provided data suitable for quantitative analyses. Although no significant differences in arthropod community diversity were found across broad geographic regions, substantial differences in community structure were observed. In North America, we also detected a decline in arthropod diversity from 10 to 5 ka BP, driven primarily by the absence of high-diversity assemblages during this period. Main Conclusions Paleomidden arthropods remain an underutilised ecological resource that can be leveraged to improve the inferential scope of arthropod research across spatiotemporal gradients. Collaboration between paleo- and neoecologists could help expand the sampling of ancient and modern arthropod communities, particularly those associated with paleomiddens, to create high-resolution datasets spanning important periods in biogeographical history. Paleomidden arthropods will remain a fundamental component for understanding arthropod biodiversity changes in the past, present, and future.

Global Ecology and Biogeography

Habitat heterogeneity, temperature, and primary productivity drive elevational gradients in avian species diversity

Aim Anticipating and mitigating the impacts of climate change on species diversity in montane ecosystems requires a mechanistic understanding of drivers of current patterns of diversity. We documented the shape of elevational gradients in avian species richness in North America and tested a suite of a priori predictions for each of five mechanistic hypotheses to explain those patterns. Location United States Methods We used predicted occupancy maps generated from species distribution models for each of 646 breeding birds to document elevational patterns in avian species richness across the six largest U.S. mountain ranges. We used spatially explicit biotic and abiotic data to test five mechanistic hypotheses proposed to explain geographic variation in species richness. Results Elevational gradients in avian species richness followed a consistent pattern of low elevation plateau-mid-elevation peak (as per McCain, 2009). We found support for three of the five hypotheses to explain the underlying cause of this pattern: the habitat heterogeneity, temperature, and primary productivity hypotheses. Main Conclusions Species richness typically decreases with elevation, but the primary cause and precise shape of the relationship remain topics of debate. We used a novel approach to study the richness-elevation relationship and our results are unique in that they show a consistent relationship between species richness and elevation among 6 mountain ranges, and universal support for three hypotheses proposed to explain the underlying cause of the observed relationship. Taken together, these results suggest that elevational variation in food availability may be the ecological process that best explains elevational gradients in avian species richness in North America. Although much attention has focused on the role of abiotic factors, particularly temperature, in limiting species’ ranges, our results offer compelling evidence that other processes also influence (and may better explain) elevational gradients in species richness.

Ecology and Evolution

Dynamic models for problems of species occurrence with multiple states

Recent extensions of occupancy modeling have focused not only on the distribution of species over space, but also on additional state variables (e.g., reproducing or not, with or without disease organisms, relative abundance categories) that provide extra information about occupied sites. These biologist-driven extensions are characterized by ambiguity in both species presence and correct state classification, caused by imperfect detection. We first show the relationships between independently published approaches to the modeling of multistate occupancy. We then extend the pattern-based modeling to the case of sampling over multiple seasons or years in order to estimate state transition probabilities associated with system dynamics. The methodology and its potential for addressing relevant ecological questions are demonstrated using both maximum likelihood (occupancy and successful reproduction dynamics of California Spotted Owl) and Markov chain Monte Carlo estimation approaches (changes in relative abundance of green frogs in Maryland). Just as multistate capture–recapture modeling has revolutionized the study of individual marked animals, we believe that multistate occupancy modeling will dramatically increase our ability to address interesting questions about ecological processes underlying population-level dynamics.

Ecology

Methods for measuring bird-mediated seed rain: Insights from a Hawaiian mesic forest

Amount and diversity of bird-dispersed seed rain play important roles in determining forest composition, yet neither is easy to quantify. The complex ecological processes that influence seed movement make the best approach highly context specific. Although recent advances in seed rain theory emphasize quantifying source-specific seed shadows, many ecological questions can be addressed u sing a less mechanistic approach that requires fewer assumptions. Using seed rain rates from 0.38 m 2 hoop traps sampled twice monthly over the course of a year, we show that number of traps required to identify changes in seed rain varies across seed species and forest type. Detecting a 50% increase in amount of seed rain required from 65 to >300 traps, while detecting a 200% increase generally required ≤⃒50 traps. Trap size and ecological context dictate the number of seeds found in each trap, but the coefficient of variation (CV) across traps in a given ecological context can help inform future studies about number of traps needed to detect change. To better understand factors influencing variation around estimates of seed rain, we simulated both clustered and evenly distributed patterns of fecal deposition using three different levels of seed aggregation (number of seeds in each fecal deposit). When patterns of fecal deposition were clustered, rather than evenly dispersed across the study area, they required >1.5 times the number of traps to identify a 100% increase in seed rain. Similarly, we found that low seed aggregation required >1.5 times the number of traps to detect a 100% change than when aggregation was medium or high. At low aggregations, fewer seed rain traps contained seeds (low, 33 ± 5%; medium, 23 ± 4%; high, 24 ± 5%), resulting in more variation across traps than medium and high aggregations. We also illustrate the importance of training observers to discern between morphologically similar seeds from different species and provide resources to help identify bird-dispersed seeds commonly found within midelevation mesic Hawaiian forests.

Hawaii

Modelling dendritic ecological networks in space: anintegrated network perspective

Dendritic ecological networks (DENs) are a unique form of ecological networks that exhibit a dendritic network topology (e.g. stream and cave networks or plant architecture). DENs have a dual spatial representation; as points within the network and as points in geographical space. Consequently, some analytical methods used to quantify relationships in other types of ecological networks, or in 2-D space, may be inadequate for studying the influence of structure and connectivity on ecological processes within DENs. We propose a conceptual taxonomy of network analysis methods that account for DEN characteristics to varying degrees and provide a synthesis of the different approaches within the context of stream ecology. Within this context, we summarise the key innovations of a new family of spatial statistical models that describe spatial relationships in DENs. Finally, we discuss how different network analyses may be combined to address more complex and novel research questions. While our main focus is streams, the taxonomy of network analyses is also relevant anywhere spatial patterns in both network and 2-D space can be used to explore the influence of multi-scale processes on biota and their habitat (e.g. plant morphology and pest infestation, or preferential migration along stream or road corridors).

Ecology Letters

Ecological effects on streams from forest fertilization: Literature review and conceptual framework for future study in the western Cascades

Fertilization of forests with urea-nitrogen has been studied numerous times for its effects on water quality. Stream nitrogen concentrations following fertilization are typically elevated during winter, including peaks in the tens-of-thousands of parts per billion range, with summer concentrations often returning to background or near-background levels. Despite these increases, water-quality criteria for nitrogen have rarely been exceeded. However, such criteria are targeted at fish toxicity or human health and are not relevant to concentrations that could cause ecological disturbances. Studies of the responses of stream biota to fertilization have been rare and have targeted either immediate, toxicity-based responses or used methods insensitive to ongoing ecological processes. This report reviews water-quality studies following forest fertilizations, emphasizing Cascade streams in the Pacific Northwest and documented biological responses in those streams. A conceptual model predicting potential ecological response to fertilization, which includes effects on algal growth and primary production, is presented. In this model, applied fertilizer nitrogen reaching streams is mostly exported during winter. However, some nitrogen retained in soils or stream and riparian areas may become available to aquatic biota during spring and summer. Biological responses may be minimal in small streams nearest to application because of light limitation, but may be elevated downstream where light is sufficient to allow algal growth. Ultimately, algal response could be greatest in downstream reaches, although ambient nutrient concentrations remain low due to uptake and benthic nutrient recycling. Ground-water flow paths and hyporheic processing could be critical in determining the fate of applied nitrogen. A framework is provided for testing this response in the Little River watershed, a tributary to the North Umpqua River, Oregon, at basic and intensive levels of investigation.

Oregon

Combined effects of climate, predation, and density dependence on Greater and Lesser Scaup population dynamics

An understanding of species relationships is critical in the management and conservation of populations facing climate change, yet few studies address how climate alters species interactions and other population drivers. We use a long-term, broad-scale data set of relative abundance to examine the influence of climate, predators, and density dependence on the population dynamics of declining scaup ( Aythya ) species within the core of their breeding range. The state-space modeling approach we use applies to a wide range of wildlife species, especially populations monitored over broad spatiotemporal extents. Using this approach, we found that immediate snow cover extent in the preceding winter and spring had the strongest effects, with increases in mean snow cover extent having a positive effect on the local surveyed abundance of scaup. The direct effects of mesopredator abundance on scaup population dynamics were weaker, but the results still indicated a potential interactive process between climate and food web dynamics (mesopredators, alternative prey, and scaup). By considering climate variables and other potential effects on population dynamics, and using a rigorous estimation framework, we provide insight into complex ecological processes for guiding conservation and policy actions aimed at mitigating and reversing the decline of scaup.

Ecological Applications

Connectivity of Mojave Desert tortoise populations—Management implications for maintaining a viable recovery network

Executive Summary The historic distribution of Mojave desert tortoises ( Gopherus agassizii ) was relatively continuous across the range, and the importance of tortoise habitat outside of designated tortoise conservation areas (TCAs) to recovery has long been recognized for its contributions to supporting gene flow between TCAs and to minimizing impacts and edge effects within TCAs. However, connectivity of Mojave desert tortoise populations has become a concern because of recent and proposed development of large tracts of desert tortoise habitat that cross, fragment, and surround designated conservation areas. This paper summarizes the underlying concepts and importance of connectivity for Mojave desert tortoise populations by reviewing current information on connectivity and providing information to managers for maintaining or enhancing desert tortoise population connectivity as they consider future proposals for development and management actions. Maintaining an ecological network for the Mojave desert tortoise, with a system of core habitats (TCAs) connected by linkages, is necessary to support demographically viable populations and long-term gene flow within and between TCAs. There are four points for wildlife and land-management agencies to consider when making decisions that could affect connectivity of Mojave desert tortoise populations (for example, in updating actions in resource management plans or amendments that could help maintain or restore functional connectivity in light of the latest information): Management of all desert tortoise habitat for persistence and connectivity . Desert tortoise populations continue to decline within most TCAs, and it is unlikely that trends are better in populations outside protected areas. Fragmentation exacerbates negative population trends by breaking large continuous populations into smaller isolated populations. Connectivity within large populations can enhance resilience to localized disturbances due to rescue by neighboring individuals. In contrast, smaller fragmented populations are resistant to rescue by their isolation and thus could suffer irreversible declines to extirpation from a variety of threats and stochastic events. Enhanced threat reduction to reverse declines within TCAs and to maintain occupied habitat in the surrounding matrix would help reduce the variability in population growth rates and improve the resilience of protected populations even while implementing efforts to improve connectivity. Each TCA has unique strengths and weaknesses regarding its ability to support minimum sustainable populations based on areal extent and its ability to support population increases based on landscape connection with adjacent populations. Considering how proposed projects (inside or outside of TCAs) affect connectivity and the ability of TCAs to support at least 5,000 adult tortoises (the numerical goal for each TCA) could help managers to maintain the resilience of TCAs to population declines. The same project, in an alternative location, could have very different impacts on local and regional populations. For example, within the habitat matrix surrounding TCAs, narrowly delineated corridors may not allow for natural population dynamics if they do not accommodate overlapping home ranges along most of their widths so that tortoises reside, grow, find mates, and produce offspring that can replace older tortoises. In addition, most habitat outside TCAs may receive more surface disturbance than habitat within TCAs. Therefore, managing the entire remaining matrix of desert tortoise habitat for permeability may be better than delineating fixed corridors. These concepts apply, especially given uncertainty about long-term condition of habitat, within and outside of TCAs under a changing climate. Ultimately, questions such as “ What are the critical linkages that need to be protected ?” could be better framed as “ How can we manage the remaining habitat matrix in ways that sustain ecological processes and habitat suitability for special status species ?” Land-management decisions made in the context of the latter question may be more conducive to maintenance of a functional ecological network. Limitations on landscape-level disturbance across habitat managed for the desert tortoise Clearly delineating habitat linkages and differentiating them from non-delineated areas by the uses that are permitted or prohibited within them by specific management guidelines can help achieve functional connectivity. Such guidelines would be most effective if they considered and accounted for all surface disturbances (for example, temporary disturbances such as fiberoptic lines or off-highway vehicle routes, right-of-ways, utility-scale solar development, urbanization) to the extent possible. A weighted framework that varies with the permanence or severity of the disturbance, and can be additive to quantify cumulative effects, could be useful (Xiong, 2020). For example, minor roads can alter tortoise movements independently of other features (Peaden and others, 2017; Hromada and others, 2020), but if the isolated dirt road is accompanied by a powerline that encourages raven predation (Xiong, 2020), then the two features together may be additive. Ignoring minor or temporary disturbance on the landscape could result in a cumulatively large impact that is not explicitly acknowledged (Goble, 2009); therefore, understanding and quantifying all surface disturbance on a given landscape is prudent. In California, the Bureau of Land Management established 0.1–1.0 percent caps on new surface-disturbance for TCAs and mapped linkages that address the issues described in number 1 of this list. Nevada, Utah, and Arizona currently do not have surface-disturbance limits. Limits comparable to those in the Desert Renewable Energy Conservation Plan (DRECP) would be 0.5 percent within TCAs and 1 percent within the linkages modeled by Averill-Murray and others (2013). Limits in some areas of California within the Desert Renewable Energy Conservation Plan, such as Ivanpah Valley, are more restrictive, at 0.1 percent. Continuity across the state line in Nevada could be achieved with comparable limits in the adjacent portion of Ivanpah Valley, as well as the Greater Trout Canyon Translocation Area and the Stump Springs Regional Augmentation Site. These more restrictive limits would help protect remaining habitat in the major interstate connectivity pathway through Ivanpah Valley and focal areas of population augmentation that provide additional population connectivity along the western flank of the Spring Mountains. In a recent study that analyzed 13 years of desert tortoise monitoring data, nearly all desert tortoise observations were at sites in which 5 percent or less of the surrounding landscape within 1 kilometer was disturbed (Carter and others, 2020a). To help maintain tortoise habitability and permeability across all other non-conservation-designated tortoise habitat, all surface disturbance could be limited to less than 5-percent development per square kilometer because the 5-percent threshold for development is the point at which tortoise occupation drops precipitously (Carter and others, 2020a). However, although individual desert tortoises were observed at development levels up to 5 percent, we do not know the fitness or reproductive characteristics of these individuals. This level of development also may not allow for long-term persistence of healthy populations that are of adequate size needed for demographic or functional connectivity; therefore, a conservative interpretation suggests that, ideally, development could be lower. Lower development levels would be particularly useful in areas within the upper 5th percentile of connectivity values modeled by Gray and others (2019). Reducing ancillary threats in places where connectivity is restricted to narrow strips of habitat, for example, narrow mountain passes or vegetated strips between solar development, could enhance the functionality of these vulnerable linkages. In such areas, maintaining multiple, redundant linkages could further enhance overall connectivity. Minimization of mortality from roads and maximization of passage under roads . Roads pose a significant threat to the long-term persistence of local tortoise populations, and roads of high traffic volume lead to severe population declines, which ultimately fragments populations farther away from the roads. Three points (a.–c.) pertain to reducing direct mortality of tortoises on the many paved roads that cross desert tortoise habitat and to maintaining a minimal level of permeability across these roads: Tortoise-exclusion fencing tied into culverts, underpasses, overpasses, or other passages below roads in desert tortoise habitat, would limit vehicular mortality of tortoises and provide opportunities for movement across the roads. Installation of shade structures on the habitat side of fences installed in areas with narrow population-depletion zones would limit overheating of tortoises that may pace the fence. Passages below highways could be maintained or retrofitted to ensure safe tortoise access, for example, by filling eroded drop-offs or modifying erosion-control features such as rip-rap to make them safer and more passable for tortoises. Wildlife management agencies could work with transportation departments to develop construction standards that are consistent with hydrologic/erosion management goals, while also incorporating a design and materials consistent with tortoise survival and passage and make the standards widely available. The process would be most effective if the status of passages was regularly monitored and built into management plans. Healthy tortoise populations along fenced highways could be supported by ensuring that land inside tortoise-exclusion fences is not so degraded that it leads to degradation of tortoise habitat outside the exclusion areas. For example, severe invasive plant infestations inside a highway exclusion could cause an increase of invasive plants outside the exclusion area and degrade habitat; therefore, invasive plants inside road rights of way could be mown or treated with herbicide to limit their spread into adjacent tortoise habitat and minimize the risk of these plants carrying wildfires into adjacent habitat. Adaptation of management based on new information . Future research will continue to build upon and refine models related to desert tortoise population connectivity and develop new ones. New models could consider landscape levels of development and be constructed such that they share common foundations to support future synthesis efforts. If model development was undertaken in partnership with entities that are responsible for management of desert tortoise habitat, it would facilitate incorporation of current and future modeling results into their land management decisions. There are specific topics that may be clarified with further evaluation: The effects of climate change on desert tortoise habitat, distribution, and population connectivity; The effects of large-scale fires, especially within repeatedly burned habitat, on desert tortoise distribution and population connectivity; The ability of solar energy facilities or similar developments to support tortoise movement and presence by leaving washes intact; leaving native vegetation intact whenever possible, or if not possible, mowing the site, allowing vegetation to re-sprout, and managing weeds; and allowing tortoises to occupy the sites; and The design and frequency of underpasses necessary to maintain functional demographic and genetic connectivity across linear features, like highways.

Arizona, California, Nevada

From theory to experiments for testing the proximate mechanisms of mast seeding: An agenda for an experimental ecology

Highly variable and synchronised production of seeds by plant populations is called masting and is implicated in many important ecological processes, but how it arises remains poorly understood. The lack of experimental studies prevents underlying mechanisms from being explicitly tested, and thereby precludes meaningful predictions on the consequences of changing environments for plant reproductive patterns and global vegetation dynamics. Here we review the most relevant hypothetical drivers of masting and outline a research agenda that takes the biology of masting from a largely observational field of ecology to one rooted in mechanistic understanding. We divide the experimental framework into three main processes: resource dynamics, pollen limitation, and genetic and hormonal regulation, and illustrate how specific predictions about proximate mechanisms can be tested, highlighting the few successful experiments as examples. We envision that the experiments we outline will deliver new insights into how and why masting patterns might respond to a changing environment.

Ecology Letters