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Understanding protected area resilience: a multi-scale, social-ecological approach

Protected areas (PAs) remain central to the conservation of biodiversity. Classical PAs were conceived as areas that would be set aside to maintain a natural state with minimal human influence. However, global environmental change and growing cross-scale anthropogenic influences mean that PAs can no longer be thought of as ecological islands that function independently of the broader social-ecological system in which they are located. For PAs to be resilient (and to contribute to broader social-ecological resilience), they must be able to adapt to changing social and ecological conditions over time in a way that supports the long-term persistence of populations, communities, and ecosystems of conservation concern. We extend Ostrom's social-ecological systems framework to consider the long-term persistence of PAs, as a form of land use embedded in social-ecological systems, with important cross-scale feedbacks. Most notably, we highlight the cross-scale influences and feedbacks on PAs that exist from the local to the global scale, contextualizing PAs within multi-scale social-ecological functional landscapes. Such functional landscapes are integral to understand and manage individual PAs for long-term sustainability. We illustrate our conceptual contribution with three case studies that highlight cross-scale feedbacks and social-ecological interactions in the functioning of PAs and in relation to regional resilience. Our analysis suggests that while ecological, economic, and social processes are often directly relevant to PAs at finer scales, at broader scales, the dominant processes that shape and alter PA resilience are primarily social and economic.

Ecological Applications

Shift of potential natural vegetation against global climate change under historical, current and future scenarios

Potential natural vegetation (PNV), the final successional stage of vegetation, plays a key role in ecological restoration, the design of nature reserves, and development of agriculture and livestock production. Meteorological data from historical and current periods including the last inter-glacial (LIG), last glacial maximum (LGM), mid Holocene (MH) periods and the present day (PD), plus derived data from 2050 and 2070, in conjunction with the Comprehensive and Sequential Classification System (CSCS) model, were used to classify global PNV. The 42 classes of global PNV were regrouped into 10 groups to facilitate analysis of spatial changes. Finally, spatio-temporal patterns and successional processes of global PNV as well as the response to climate changes were analysed. Our study made the following five conclusions. (1) Only one missing class (IA1 frigid-extrarid frigid desert, alpine desert) arose in periods of LIG, MH, 2050, and 2070 for global PNV. (2) The frigid-arid groups were mainly distributed in higher latitudes and elevations, but temperate-humid groups and tropical-perhumid groups occurred in middle and low latitudes, respectively. Temperate zonal forest steppe, warm desert, savanna and tropical zonal forest steppe increased, while six other groups decreased. (3) The conversion from temperate zonal forest steppe to tundra and alpine steppe from LIG to LGM occupied the largest area, indicating a drastic shift in climate and the associated response of terrestrial vegetation sensitive to climate change. (4) The CSCS could be used to simulate the long-term succession of global PNV. (5) As a consequence of global warming, forests shifted to the northern hemisphere and Tibet, areas with much higher latitude and elevation. The PNV groups with greater shift distance revealed the more serious effects of global climate change on vegetation.

Rangeland Journal

Biological soil crusts (biocrusts) as a model system in community, landscape and ecosystem ecology

Model systems have had a profound influence on the development of ecological theory and general principles. Compared to alternatives, the most effective models share some combination of the following characteristics: simpler, smaller, faster, general, idiosyncratic or manipulable. We argue that biological soil crusts (biocrusts) have unique combinations of these features that should be more widely exploited in community, landscape and ecosystem ecology. In community ecology, biocrusts are elucidating the importance of biodiversity and spatial pattern for maintaining ecosystem multifunctionality due to their manipulability in experiments. Due to idiosyncrasies in their modes of facilitation and competition, biocrusts have led to new models on the interplay between environmental stress and biotic interactions and on the maintenance of biodiversity by competitive processes. Biocrusts are perhaps one of the best examples of micro-landscapes—real landscapes that are small in size. Although they exhibit varying patch heterogeneity, aggregation, connectivity and fragmentation, like macro-landscapes, they are also compatible with well-replicated experiments (unlike macro-landscapes). In ecosystem ecology, a number of studies are imposing small-scale, low cost manipulations of global change or state factors in biocrust micro-landscapes. The versatility of biocrusts to inform such disparate lines of inquiry suggests that they are an especially useful model system that can enable researchers to see ecological principles more clearly and quickly.

Biodiversity and Conservation

Effects of temperature on disease progression and swimming stamina in Ichthyophonus‐infected rainbow trout, Oncorhynchus mykiss (Walbaum)

Rainbow trout, Oncorhynchus mykiss , were infected with Ichthyophonus sp. and held at 10°C, 15°C and 20°C for 28 days to monitor mortality and disease progression. Infected fish demonstrated more rapid onset of disease, higher parasite load, more severe host tissue reaction and reduced mean‐day‐to‐death at higher temperature. In a second experiment, Ichthyophonus ‐infected fish were reared at 15°C for 16 weeks then subjected to forced swimming at 10°C, 15°C and 20°C. Stamina improved significantly with increased temperature in uninfected fish; however, this was not observed for infected fish. The difference in performance between infected and uninfected fish became significant at 15°C ( P =0.02) and highly significant at 20°C ( P = 0.005). These results have implications for changes in the ecology of fish diseases in the face of global warming and demonstrate the effects of higher temperature on the progression and severity of ichthyophoniasis as well as on swimming stamina, a critical fitness trait of salmonids. This study helps explain field observations showing the recent emergence of clinical ichthyophoniasis in Yukon River Chinook salmon later in their spawning migration when water temperatures were high, as well as the apparent failure of a substantial percentage of infected fish to successfully reach their natal spawning areas.

Journal of Fish Diseases

Sprague’s Pipits (Anthus spragueii) occupying high-elevation intermontane valley habitat throughout the breeding season in southwest Montana

Sprague's Pipit ( Anthus spragueii ) is an imperiled bird species that breeds in grasslands of the northern Great Plains of North America. While conducting avian surveys in the Centennial Valley, Beaverhead County, southwest Montana, during spring and summer 2023, we detected approximately 10 male Sprague's Pipits performing aerial displays. These represent the first detailed observations of Sprague's Pipits in Beaverhead County, Montana, and the first confirmed observations of territorial displays therein. Subsequent detections included multiple adults, one of which was seen flying with food in its bill. We hypothesize that these observations represent a small but previously undescribed breeding population occurring above 2000 m elevation, which is higher than elevations at which this species is generally thought to breed. Our discovery also expands the known breeding habitat of this species to include intermontane valley bottoms, raising questions of whether the historical range included this habitat prior to its large-scale modifications across western North America and contemporary global change. These additions to our understanding of Sprague's Pipit ecology may be important for both land managers in the region and conservation efforts for this species. The evidence described here, of a breeding population of Sprague's Pipits in the Centennial Valley, Montana, may warrant further investigations in intermontane valleys throughout the region to better understand the breeding extent and the management actions needed to conserve this species and its habitat.

Montana

Ecological prediction at macroscales using big data: Does sampling design matter?

Although ecosystems respond to global change at regional to continental scales (i.e., macroscales), model predictions of ecosystem responses often rely on data from targeted monitoring of a small proportion of sampled ecosystems within a particular geographic area. In this study, we examined how the sampling strategy used to collect data for such models influences predictive performance. We subsampled a large and spatially-extensive dataset to investigate how macroscale sampling strategy affects prediction of ecosystem characteristics in 6,784 lakes across a 1.8 million km2 area. We estimated model predictive performance for different subsets of the dataset to mimic three common sampling strategies for collecting observations of ecosystem characteristics: random sampling design, stratified random sampling design, and targeted sampling. We found that sampling strategy influenced model predictive performance such that (1) stratified random sampling designs did not improve predictive performance compared to simple random sampling designs and (2) although one of the scenarios that mimicked targeted (non-random) sampling had the poorest performing predictive models, the other targeted sampling scenarios resulted in models with similar predictive performance to that of the random sampling scenarios. Our results suggest that although potential biases in datasets from some forms of targeted sampling may limit predictive performance, compiling existing spatially-extensive datasets can result in models with good predictive performance that may inform a wide range of science questions and policy goals related to global change.

Ecological Applications

Three decades of World Recreational Fishing Conferences: What have we learned about the dynamics of recreational fisheries?

Recreational fishing extends beyond catching fish, connecting individuals with nature, generating environmental stewards and contributing significantly to both local, regional and national economies. Assessing potential impacts on ecosystems and social-ecological systems requires dedicated multi- and interdisciplinary research and integrative management and policy approaches. Managing recreational fisheries effectively also necessitates transdisciplinary thinking, yet this poses continued challenges, as researchers and managers balance specialized expertise with innovative, boundary-crossing perspectives in light of limited funding in a rapidly changing social-ecological environment. The World Recreational Fishing Conference (WRFC) serves as the leading global scientific forum for addressing these challenges, fostering interdisciplinary exchange among scientists, managers, policy makers and stakeholders. The papers in this special issue represent the output of the last WRFC in Melbourne, Australia in 2023. By reviewing the content of the papers published in the present special issue, we illustrate the critical role of collaborative spaces, such as the WRFC, in bridging different expertise and approaches and fostering innovation, thereby securing adaptive management and conservation of recreational fisheries in response to global changes. Evaluating the development of the WRFC over three decades highlights how this conference series represents a leading think tank serving the entire network of recreational fisheries professionals globally.

Fisheries Management and Ecology

The distribution and role of functional abundance in cross‐scale resilience

The cross‐scale resilience model suggests that system‐level ecological resilience emerges from the distribution of species’ functions within and across the spatial and temporal scales of a system. It has provided a quantitative method for calculating the resilience of a given system and so has been a valuable contribution to a largely qualitative field. As it is currently laid out, the model accounts for the spatial and temporal scales at which environmental resources and species are present and the functional roles species play but does not inform us about how much resource is present or how much function is provided. In short, it does not account for abundance in the distribution of species and their functional roles within and across the scales of a system. We detail the ways in which we would expect species’ abundance to be relevant to the cross‐scale resilience model based on the extensive abundance literature in ecology. We also put forward a series of testable hypotheses that would improve our ability to anticipate and quantify how resilience is generated, and how ecosystems will (or will not) buffer recent rapid global changes. This stream of research may provide an improved foundation for the quantitative evaluation of ecological resilience.

Ecology

The role of climate in the dynamics of a hybrid zone in Appalachian salamanders

I examined the potential influence of climate change on the dynamics of a previously studied hybrid zone between a pair of terrestrial salamanders at the Coweeta Hydrologic Laboratory, U.S. Forest Service, in the Nantahala Mountains of North Carolina, USA. A 16-year study led by Nelson G. Hairston, Sr. revealed that Plethodon teyahalee and Plethodon shermani hybridized at intermediate elevations, forming a cline between 'pure' parental P. teyahalee at lower elevations and 'pure' parental P. shermani at higher elevations. From 1974 to 1990 the proportion of salamanders at the higher elevation scored as 'pure' P. shermani declined significantly, indicating that the hybrid zone was spreading upward. To date there have been no rigorous tests of hypotheses for the movement of this hybrid zone. Using temperature and precipitation data from Coweeta, I re-analyzed Hairston's data to examine whether the observed elevational shift was correlated with variation in either air temperature or precipitation from the same time period. For temperature, my analysis tracked the results of the original study: the proportion of 'pure' P. shermani at the higher elevation declined significantly with increasing mean annual temperature, whereas the proportion of 'pure' P. teyahalee at lower elevations did not. There was no discernable relationship between proportions of 'pure' individuals of either species with variation in precipitation. From 1974 to 1990, low-elevation air temperatures at the Coweeta Laboratory ranged from annual means of 11.8 to 14.2 °C, compared with a 55-year average (1936-1990) of 12.6 °C. My re-analyses indicate that the upward spread of the hybrid zone is correlated with increasing air temperatures, but not precipitation, and provide an empirical test of a hypothesis for one factor that may have influenced this movement. My results aid in understanding the potential impact that climate change may have on the ecology and evolution of terrestrial salamanders in montane regions.

Global Change Biology

Where has turtle ecology been, and where is it going?

Over 9000 articles have been published on turtles and tortoises (excluding sea turtles) since 1950 according to the Web of Science, including over 8000 contained in a personal bibliography that we analyze in this paper. Research had a slow start from 1900 to 1950, with mostly anecdotal additions to our knowledge until the contributions of F. Cagle and A. Carr took turtle research to new levels as the cofathers of turtle ecology in the middle of the last century. Books written in 1939, 1952, and 1972 that compiled existing literature on turtles in the United States and Canada set the stage for growing interest in turtles. The first global compilation of turtle species did not become available until 1961. Publication frequency increased in the 1960s and especially the 1970s as interest in turtles grew, and a wave of turtle biologists emerged from doctoral degree programs. We briefly review the contributions of scientists who published extensively on turtle ecology in those and later decades up to the present. We also review advances in our knowledge of various topics, including the global distribution of turtle research efforts; changes in our perceptions of turtle species diversity over time; turtle community ecology; sex ratios, sex-determination, and climate change; overwintering behavior; sexual size dimorphism and sexual dichromatism; analyses of population genetics; turtles and vocalization; and the emergence of turtle conservation biology efforts. We conclude with a discussion of future opportunities and challenges for working with turtles.

Herpetologica

Are we heading towards a global decrease in coregonine catches?

Coregonines have specific ecological needs, making them potentially very vulnerable to changes in lake conditions. A contemporary concern is that many lakes worldwide are experiencing environmental changes due to anthropogenic pressure and climate warming. Here, we compiled long-term data of coregonine catches from 27 lakes from three continents in the northern hemisphere. Declines in catch were observed in 67% of the lakes during the first two decades of the 21 st century, with a significant trend (p-value<0.05) in 44% of the cases. An analysis to determine whether trends are globally linked to environmental conditions and specific lake attributes was carried out on 26 lakes for the period 2000–2019. Several local declines in catches had already been documented in the literature and are likely to be due to local forcing such as nutrients, species invasions and changes in fishing practices. Nevertheless, on a global scale, our results indicate that lakes, which exhibited a significant decrease in catch were larger and more nutrient-poor than other lakes. The rate of change in catches appeared to be related to the trophic state of the lake. The specific effect of warming is difficult to determine during the studied period. When warming occurred outside the period of egg incubation, decreasing trends were more frequently observed in nutrient-poor than in mesotrophic lakes. In conclusion, our findings suggest that achieving oligo- or ultraoligotrophic conditions, as required in developed countries to control phytoplankton blooms and enhance water quality, could pose significant challenges for future management of coregonine fisheries.

International Journal of Limnology

Soil cover heterogeneity associated with biocrusts predicts patch-level plant diversity patterns

Context Soil resource heterogeneity drives plant species diversity patterns at local and landscape scales. In drylands, biocrusts are patchily distributed and contribute to soil resource heterogeneity important for plant establishment and growth. Yet, we have a limited understanding of how such heterogeneity may relate to patterns of plant diversity and community structure. Objectives We explored relationships between biocrust-associated soil cover heterogeneity and plant diversity patterns in a cool desert ecosystem. We asked: (1) does biocrust-associated soil cover heterogeneity predict plant diversity and community composition? and (2) can we use high-resolution remote sensing data to calculate soil cover heterogeneity metrics that could be used to extrapolate these patterns across landscapes? Methods We tested associations among field-based measures of plant diversity and soil cover heterogeneity. We then used a Support Vector Machine classification to map soil, plant and biocrust cover from sub-centimeter resolution Unoccupied Aerial System (UAS) imagery and compared the mapped results to field-based measures. Results Field-based soil cover heterogeneity and biocrust cover were positively associated with plant diversity and predicted community composition. The accuracy of UAS-mapped soil cover classes varied across sites due to variation in timing and quality of image collections, but the overall results suggest that UAS are a promising data source for generating detailed, spatially explicit soil cover heterogeneity metrics. Conclusions Results improve understanding of relationships between biocrust-associated soil cover heterogeneity and plant diversity and highlight the promise of high-resolution UAS data to extrapolate these patterns over larger landscapes which could improve conservation planning and predictions of dryland responses to soil degradation under global change.

Utah

Groundwater declines are linked to changes in Great Plains stream fish assemblages

Groundwater pumping for agriculture is a major driver causing declines of global freshwater ecosystems, yet the ecological consequences for stream fish assemblages are rarely quantified. We combined retrospective (1950–2010) and prospective (2011–2060) modeling approaches within a multiscale framework to predict change in Great Plains stream fish assemblages associated with groundwater pumping from the United States High Plains Aquifer. We modeled the relationship between the length of stream receiving water from the High Plains Aquifer and the occurrence of fishes characteristic of small and large streams in the western Great Plains at a regional scale and for six subwatersheds nested within the region. Water development at the regional scale was associated with construction of 154 barriers that fragment stream habitats, increased depth to groundwater and loss of 558 km of stream, and transformation of fish assemblage structure from dominance by large-stream to small-stream fishes. Scaling down to subwatersheds revealed consistent transformations in fish assemblage structure among western subwatersheds with increasing depths to groundwater. Although transformations occurred in the absence of barriers, barriers along mainstem rivers isolate depauperate western fish assemblages from relatively intact eastern fish assemblages. Projections to 2060 indicate loss of an additional 286 km of stream across the region, as well as continued replacement of large-stream fishes by small-stream fishes where groundwater pumping has increased depth to groundwater. Our work illustrates the shrinking of streams and homogenization of Great Plains stream fish assemblages related to groundwater pumping, and we predict similar transformations worldwide where local and regional aquifer depletions occur.

Colorado, Kansas, Nebraska

Applying the ecology of aquatic–terrestrial linkages to freshwater and riparian management

Global stressors such as climate change, invasive species, urbanization, agricultural practices, and pollution can alter aquatic resource subsidies to terrestrial consumers. The effects of these stressors on timing, quality, and quantity of aquatic subsidies, such as adult aquatic insects, to birds, herpetofauna, and mammals, have large implications for wildlife management (Baxter et al. 2004 , Saunders and Fausch 2007 , Walters et al. 2008 , Sullivan and Rodewald 2012 , Morrissey et al. 2015 , Kraus et al. 2016 , Larsen et al. 2016 , Sullivan et al. 2019 ). For example, insect-mediated contaminant transport from polluted rivers expose song bird nestlings and other protected birds to potentially toxic levels of persistent organic contaminants and pharmaceuticals (Walters et al. 2010 , Richmond et al. 2018 ). Recent declines in aquatic insect production caused by pollution and changes in land use have been tied to global declines in terrestrial insectivores such as birds and bats (Hallmann et al. 2014 , Morrissey et al. 2015 , Raby et al. 2018 ). These natural-resource impacts from aquatic–terrestrial exposure and loss of resource subsidies are leading to changes in monitoring protocols and how management agencies evaluate the effectiveness of corrective remedies (Muehlbauer et al. 2019 ). For researchers interested in the application of resource subsidy research, a logical next step is to help practitioners anticipate effects of global stressors on aquatic-terrestrial linkages and incorporate these principles into decision making.

Freshwater Science

Impacts of climatic variation on trout: A global synthesis and path forward

Despite increasing concern that climate change may negatively impact trout&mdash;a globally distributed group of fish with major economic, ecological, and cultural value&mdash;a synthetic assessment of empirical data quantifying relationships between climatic variation and trout ecology does not exist. We conducted a systematic review to describe how temporal variation in temperature and streamflow influences trout ecology in freshwater ecosystems. Few studies ( n = 42) have quantified relationships between temperature or streamflow and trout demography, growth, or phenology, and nearly all estimates (96 %) were for Salvelinus fontinalis and Salmo trutta . Only seven studies used temporal data to quantify climate-driven changes in trout ecology. Results from these studies were beset with limitations that prohibited quantitatively rigorous meta-analysis, a concerning inadequacy given major investment in trout conservation and management worldwide. Nevertheless, consistent patterns emerged from our synthesis, particularly a positive effect of summer streamflow on trout demography and growth; 64 % of estimates were positive and significant across studies, age classes, species, and locations, highlighting that climate-induced changes in hydrology may have numerous consequences for trout. To a lesser degree, summer and fall temperatures were negatively related to population demography (51 and 53 % of estimates, respectively), but temperature was rarely related to growth. To address limitations and uncertainties, we recommend: (1) systematically improving data collection, description, and sharing; (2) appropriately integrating climate impacts with other intrinsic and extrinsic drivers over the entire lifecycle; (3) describing indirect consequences of climate change; and (4) acknowledging and describing intrinsic resiliency.

Reviews in Fish Biology and Fisheries

A river basin spatial model to quantitively advance understanding of riverine tree response dynamics to water availability and hydrological management

Ecological condition continues to decline in arid and semi-arid river basins globally due to hydrological over-abstraction combined with changing climatic conditions. Whilst provision of water for the environment has been a primary approach to alleviate ecological decline, how to accurately monitor changes in riverine trees at fine spatial and temporal scales, remains a substantial challenge. This is further complicated by constantly changing water availability across expansive river basins with varying climatic zones. Within, we combine rare, fine-scale, high frequency temporal in-situ field collected data with machine learning and remote sensing, to provide a robust model that enables broadscale monitoring of physiological tree water stress response to environmental changes via actual evapotranspiration (ET). Physiological variation of Eucalyptus camaldulensis (River Red Gum) and E. largiflorens (Black Box) trees across 10 study locations in the southern Murray-Darling Basin, Australia, was captured instantaneously using sap flow sensors, substantially reducing tree response lags encountered by monitoring visual canopy changes. Actual ET measurement of both species was used to bias correct a national spatial ET product where a Random Forest model was trained using continuous timeseries of in-situ data of up to four years. Precise monthly AMLETT ( A ustralia-wide M achine L earning ET for T rees) ET outputs in 30 m pixel resolution from 2012 to 2021, were derived by incorporating additional remote sensing layers such as soil moisture, land surface temperature, radiation and EVI and NDVI in the Random Forest model. Landsat and Sentinal-2 correlation results between in-situ ET and AMLETT ET returned R 2 of 0.94 (RMSE 6.63 mm period −1 ) and 0.92 (RMSE 6.89 mm period −1 ), respectively. In comparison, correlation between in-situ ET and a national ET product returned R 2 of 0.44 (RMSE 34.08 mm period −1 ) highlighting the need for bias correction to generate accurate absolute ET values. The AMLETT method presented here, enhances environmental management in river basins worldwide. Such robust broadscale monitoring can inform water accounting and importantly, assist decisions on where to prioritize water for the environment to restore and protect key ecological assets and preserve floodplain and riparian ecological function.

The Journal of Environmental Management

The human dimension of fire regimes on Earth

Humans and their ancestors are unique in being a fire‐making species, but ‘natural’ (i.e. independent of humans) fires have an ancient, geological history on Earth. Natural fires have influenced biological evolution and global biogeochemical cycles, making fire integral to the functioning of some biomes. Globally, debate rages about the impact on ecosystems of prehistoric human‐set fires, with views ranging from catastrophic to negligible. Understanding of the diversity of human fire regimes on Earth in the past, present and future remains rudimentary. It remains uncertain how humans have caused a departure from ‘natural’ background levels that vary with climate change. Available evidence shows that modern humans can increase or decrease background levels of natural fire activity by clearing forests, promoting grazing, dispersing plants, altering ignition patterns and actively suppressing fires, thereby causing substantial ecosystem changes and loss of biodiversity. Some of these contemporary fire regimes cause substantial economic disruptions owing to the destruction of infrastructure, degradation of ecosystem services, loss of life, and smoke‐related health effects. These episodic disasters help frame negative public attitudes towards landscape fires, despite the need for burning to sustain some ecosystems. Greenhouse gas‐induced warming and changes in the hydrological cycle may increase the occurrence of large, severe fires, with potentially significant feedbacks to the Earth system. Improved understanding of human fire regimes demands: (1) better data on past and current human influences on fire regimes to enable global comparative analyses, (2) a greater understanding of different cultural traditions of landscape burning and their positive and negative social, economic and ecological effects, and (3) more realistic representations of anthropogenic fire in global vegetation and climate change models. We provide an historical framework to promote understanding of the development and diversification of fire regimes, covering the pre‐human period, human domestication of fire, and the subsequent transition from subsistence agriculture to industrial economies. All of these phases still occur on Earth, providing opportunities for comparative research.

Journal of Biogeography

A mosaic-based approach is needed to conserve biodiversity in disturbed freshwater ecosystems

Conserving native biodiversity in the face of human‐ and climate‐related impacts is a challenging and globally important ecological problem that requires an understanding of spatially connected, organismal‐habitat relationships. Globally, a suite of disturbances (e.g., agriculture, urbanization, climate change) degrades habitats and threatens biodiversity. A mosaic approach (in which connected, interacting collections of juxtaposed habitat patches are examined) provides a scientific foundation for addressing many disturbance‐related, ecologically based conservation problems. For example, if specific habitat types disproportionately increase biodiversity, these keystones should be incorporated into research and management plans. Our sampling of fish biodiversity and aquatic habitat along ten 3‐km sites within the Upper Neosho River subdrainage, KS, from June‐August 2013 yielded three generalizable ecological insights. First, specific types of mesohabitat patches (i.e., pool, riffle, run, and glide) were physically distinct and created unique mosaics of mesohabitats that varied across sites. Second, species richness was higher in riffle mesohabitats when mesohabitat size reflected field availability. Furthermore, habitat mosaics that included more riffles had greater habitat diversity and more fish species. Thus, riffles (<5% of sampled area) acted as keystone habitats. Third, additional conceptual development, which we initiate here, can broaden the identification of keystone habitats across ecosystems and further operationalize this concept for research and conservation. Thus, adopting a mosaic approach can increase scientific understanding of organismal‐habitat relationships, maintain natural biodiversity, advance spatial ecology, and facilitate effective conservation of native biodiversity in human‐altered ecosystems.

Kansas