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Stoneflies in the genus Lednia (Plecoptera: Nemouridae): Sentinels of climate change impacts on mountain stream biodiversity

Rapid recession of glaciers and snowfields is threatening the habitats of cold-water biodiversity worldwide. In many ice-sourced headwaters of western North America, stoneflies in the genus Lednia (Plecoptera: Nemouridae) are a prominent member of the invertebrate community. With a broad distribution in mountain streams and close ties to declining glacier cover, Lednia has emerged as a sentinel of climate change threats to high-elevation aquatic biodiversity. Lednia tumana , which is endemic to Glacier National Park, USA and the surrounding mountains, is the most well-studied species in the genus and in 2019 became federally protected under the U.S. Endangered Species Act (ESA) due to climate-induced loss of meltwater habitats. Three other Lednia species have also been described, and like L. tumana , each is endemic to a mountain region of western North America: Lednia sierra in the Sierra Nevada, Lednia borealis in the Cascade Range, and Lednia tetonica in the Teton Range. In this review, we provide a comprehensive overview of Lednia ecology, genetics, and physiology, with an emphasis on the conservation outlook for the group and species with similar headwater distributions. We highlight substantial progress made in the last decade to better understand the ecology and evolution of Lednia , including the identification of 140 Lednia- containing streams (an increase from 12 streams in 2010), and a more complete understanding of the degree to which warming streams may imperil species in the genus. In light of the ESA listing of L. tumana , we show that similar conservation threats likely face all extant Lednia species. However, substantial gaps in our knowledge remain, primarily centering around their distributions (and the potential for as yet undescribed species), life history, ecophysiology, and trophic ecology. We conclude by describing pressing questions for Lednia that when addressed will expand knowledge of the genus and its conservation as well as broader understanding of climate risks to mountain stream biodiversity worldwide.

California, Montana, Washington, Wyoming

Mediterranean biomes: Evolution of their vegetation, floras and climate

Mediterranean-type ecosystems (MTEs) possess the highest levels of plant species richness in the world outside of the wet tropics. Sclerophyll vegetation similar to today’s mediterranean-type shrublands was already present on oligotrophic soils in the wet and humid climate of the Cretaceous, with fire-adapted Paleogene lineages in southwestern Australia and the Cape Region. The novel MTC seasonality present since the mid-Miocene has allowed colonization of MTEs from a regional species pool with associated diversification. Fire persistence has been a primary driving factor for speciation in four of the five regions. Understanding the regional patterns of plant species diversity among the MTEs involves complex interactions of geologic and climatic histories for each region as well as ecological factors that have promoted diversification in the Neogene and Quaternary. A critical element of species richness for many MTE lineages has been their ability to speciate and persist at fine spatial scales, with low rates of extinction.

Annual Review of Ecology, Evolution, and Systemati

Migratory flyway and geographical distance are barriers to the gene flow of influenza virus among North American birds

Despite the importance of migratory birds in the ecology and evolution of avian influenza virus (AIV), there is a lack of information on the patterns of AIV spread at the intra‐continental scale. We applied a variety of statistical phylogeographic techniques to a plethora of viral genome sequence data to determine the strength, pattern and determinants of gene flow in AIV sampled from wild birds in North America. These analyses revealed a clear isolation‐by‐distance of AIV among sampling localities. In addition, we show that phylogeographic models incorporating information on the avian flyway of sampling proved a better fit to the observed sequence data than those specifying homogeneous or random rates of gene flow among localities. In sum, these data strongly suggest that the intra‐continental spread of AIV by migratory birds is subject to major ecological barriers, including spatial distance and avian flyway.

California, Oregon, Washington

Integrating Earth–life systems: A geogenomic approach

For centuries, scientists have recognized and worked to understand how Earth’s mutable landscape and climate shape the distribution and evolution of species. Here, we describe the emerging field of geogenomics, which uses the reciprocal and deep integration of geologic, climatic, and population genomic data to define and test cause–effect relationships between Earth and life at intermediate spatial and temporal scales (i.e., the mesoscale). Technological advances now power the detailed reconstruction of landscape and evolutionary histories, but transdisciplinary collaborations and new quantitative tools are needed to better integrate Earth–life data. Geogenomics can help build a more unified theory and characterize the boundary conditions under which geologic and climatic processes generate new biodiversity, how species’ responses differ, and why.

Trends in Ecology & Evolution

Paleoclimate ocean conditions shaped the evolution of corals and their skeletal composition through deep time

Identifying how past environmental conditions shaped the evolution of corals and their skeletal traits provides a framework for predicting their persistence and that of their non-calcifying relatives under impending global warming and ocean acidification. Here we show that ocean geochemistry, particularly aragonite–calcite seas, drives patterns of morphological evolution in anthozoans (corals, sea anemones) by examining skeletal traits in the context of a robust, time-calibrated phylogeny. The lability of skeletal composition among octocorals suggests a greater ability to adapt to changes in ocean chemistry compared with the homogeneity of the aragonitic skeleton of scleractinian corals. Pulses of diversification in anthozoans follow mass extinctions and reef crises, with sea anemones and proteinaceous corals filling empty niches as tropical reef builders went extinct. Changing environmental conditions will likely diminish aragonitic reef-building scleractinians, but the evolutionary history of the Anthozoa suggests other groups will persist and diversify in their wake.

Nature Ecology & Evolution

Geochemistry of the Black Sea during the last 15 kyr: A protracted evolution of its hydrography and ecology

The Black Sea is a 2200 m deep anoxic, marine sea connected to the Mediterranean Sea via the Dardanelles Strait, Marmara Sea, and the 3 km wide, 35 m deep Bosphorus Strait. The biogeochemistry of sediment from the Anatolia slope has recorded changes to the hydrography leading up to and following the input of Mediterranean water at ~9.4 ka (10 3 years B.P.), when global sea level rose to the level of the Bosphorus sill and high-salinity water from the Mediterranean began to spill into the then brackish lake. The water initially mixed little with the lake water but cascaded to the bottom where it remained essentially isolated for ~1.6 kyr, the time required to fill the basin from the bottom up at its present input rate. The accumulation of Mo in the seafloor sediments, a proxy of bottom-water anoxia, increased sharply at ~8.6 ka, when bacterial respiration in the bottom water advanced to SO 4 2− reduction by the oxidation of organic detritus that settled out of the photic zone. Its accumulation remained elevated to ~5.6 ka, when it decreased 60%, only to again increase slightly at ~2.0 ka. The accumulation of C org , a proxy of primary productivity, increased threefold to fourfold at ~7.8 ka, when upward mixing of the high-salinity bottom water replaced the then thin veneer of the brackish photic zone in less than 50 years. From that time onward, the accumulation of C org , Mo, and additional trace metals has reflected the hydrography of the basin and Bosphorus Strait, controlled largely by climate.

Paleoceanography

Spatial sampling bias and model complexity in stream-based species distribution models: A case study of Paddlefish (Polyodon spathula) in the Arkansas River basin, USA

Leveraging existing presence records and geospatial datasets, species distribution modeling has been widely applied to informing species conservation and restoration efforts. Maxent is one of the most popular modeling algorithms, yet recent research has demonstrated Maxent models are vulnerable to prediction errors related to spatial sampling bias and model complexity. Despite elevated rates of biodiversity imperilment in stream ecosystems, the application of Maxent models to stream networks has lagged, as has the availability of tools to address potential sources of error and calculate model evaluation metrics when modeling in nonraster environments (such as stream networks). Herein, we use Maxent and customized R code to estimate the potential distribution of paddlefish ( Polyodon spathula ) at a stream-segment level within the Arkansas River basin, USA, while accounting for potential spatial sampling bias and model complexity. Filtering the presence data appeared to adequately remove an eastward, large-river sampling bias that was evident within the unfiltered presence dataset. In particular, our novel riverscape filter provided a repeatable means of obtaining a relatively even coverage of presence data among watersheds and streams of varying sizes. The greatest differences in estimated distributions were observed among models constructed with default versus AIC C -selected parameterization. Although all models had similarly high performance and evaluation metrics, the AIC C -selected models were more inclusive of westward-situated and smaller, headwater streams. Overall, our results solidified the importance of accounting for model complexity and spatial sampling bias in SDMs constructed within stream networks and provided a roadmap for future paddlefish restoration efforts in the study area.

Arkansas, Colorado, Kansas, Missouri, Nebraska, Ne

The cost of self-defense: Browsing effects in the rare plant species Salix arizonica

Coevolution between plants and their animal predators has led to diverse defensive adaptations. Multiple theories of defense propose that there are resource allocation costs associated with producing chemical defenses. One leading hypothesis, optimal defense theory (ODT), suggests that natural selection will result in the allocation of resources to defenses that optimize the cost-to-benefit ratio between defense and other functional processes. The population decline of the rare subalpine wetland species, Arizona willow ( Salix arizonica ), has been attributed to various biotic and abiotic factors, with browsing from wild and domestic ungulates as a significant concern for at least three decades. In a field experiment using natural populations, we compare the relationship between phytochemical defense and height in Arizona willows with and without long-term protection from browsing via browse exclosures. Consistent with the predictions of ODT, individuals with physical protection from ungulate browsing for multiple years had significantly lower phenolic glycoside (PG) concentrations and increased plant height compared to unprotected individuals. A similar pattern was found across all individuals, whereby total PG concentration and height were negatively correlated. In a short-term experiment in natural populations, changes in levels of defense were not observed when plants received protection for only one growing season. The contrasting pattern of defense plasticity in response to long-term versus short-term physical protection suggests a differential plastic response in this long-lived species. Delayed reduction in PG concentration may serve as a benefit to avoid mismatches between environmental cues and responses. Our research sheds light on the intricate dynamics between plant-defense strategies, environmental pressures, and evolutionary adaptations in shaping plant–browser interactions.

Arizona, Colorado, New Mexico, Utah

Spatial capture–recapture with random thinning for unidentified encounters

Spatial capture–recapture (SCR) models have increasingly been used as a basis for combining capture–recapture data types with variable levels of individual identity information to estimate population density and other demographic parameters. Recent examples are the unmarked SCR (or spatial count model), where no individual identities are available and spatial mark–resight (SMR) where individual identities are available for only a marked subset of the population. Currently lacking, though, is a model that allows unidentified samples to be combined with identified samples when there are no separate classes of “marked” and “unmarked” individuals and when the two sample types cannot be considered as arising from two independent observation models. This is a common scenario when using noninvasive sampling methods, for example, when analyzing data on identified and unidentified photographs or scats from the same sites. Here we describe a “random thinning” SCR model that utilizes encounters of both known and unknown identity samples using a natural mechanistic dependence between samples arising from a single observation model. Our model was fitted in a Bayesian framework using NIMBLE. We investigate the improvement in parameter estimates by including the unknown identity samples, which was notable (up to 79% more precise) in low‐density populations with a low rate of identified encounters. We then applied the random thinning SCR model to a noninvasive genetic sampling study of brown bear ( Ursus arctos ) density in Oriental Cantabrian Mountains (North Spain). Our model can improve density estimation for noninvasive sampling studies for low‐density populations with low rates of individual identification, by making use of available data that might otherwise be discarded.

Ecology & Evolution

Genetic and morphological characterization of the freshwater mussel clubshell species complex (Pleurobema clava and Pleurobema oviforme) to inform conservation planning

The shell morphologies of the freshwater mussel species Pleurobema clava (federally endangered) and Pleurobema oviforme (species of concern) are similar, causing considerable taxonomic confusion between the two species over the last 100 years. While P. clava was historically widespread throughout the Ohio River basin and tributaries to the lower Laurentian Great Lakes, P. oviforme was confined to the Tennessee and the upper Cumberland River basins. We used two mitochondrial DNA (mtDNA) genes, 13 novel nuclear DNA microsatellite markers, and shell morphometrics to help resolve this taxonomic confusion. Evidence for a single species was apparent in phylogenetic analyses of each mtDNA gene, revealing monophyletic relationships with minimal differentiation and shared haplotypes. Analyses of microsatellites showed significant genetic structuring, with four main genetic clusters detected, respectively, in the upper Ohio River basin, the lower Ohio River and Great Lakes, and upper Tennessee River basin, and a fourth genetic cluster, which included geographically intermediate populations in the Ohio and Tennessee river basins. While principal components analysis (PCA) of morphometric variables (i.e., length, height, width, and weight) showed significant differences in shell shape, only 3% of the variance in shell shape was explained by nominal species. Using Linear Discriminant and Random Forest (RF) analyses, correct classification rates for the two species' shell forms were 65.5% and 83.2%, respectively. Random Forest classification rates for some populations were higher; for example, for North Fork Holston (HOLS), it was >90%. While nuclear DNA and shell morphology indicate that the HOLS population is strongly differentiated, perhaps indicative of cryptic biodiversity, we consider the presence of a single widespread species the most likely biological scenario for many of the investigated populations based on our mtDNA dataset. However, additional sampling of P. oviforme populations at nuclear loci is needed to corroborate this finding.

Indiana, Kentucky, Ohio, Pennsylvania, Tennessee,

Abdominally implanted satellite transmitters affect reproduction and survival rather than migration of large shorebirds

Satellite telemetry has become a common technique to investigate avian life-histories, but whether such tagging will affect fitness is a critical unknown. In this study, we evaluate multi-year effects of implanted transmitters on migratory timing and reproductive performance in shorebirds. Shorebirds increasingly are recognized as good models in ecology and evolution. That many of them are of conservation concern adds to the research responsibilities. In May 2009, we captured 56 female Black-tailed Godwits Limosa limosa limosa during late incubation in The Netherlands. Of these, 15 birds were equipped with 26-g satellite transmitters with a percutaneous antenna (7.8 % ± 0.2 SD of body mass), surgically implanted in the coelom. We compared immediate nest survival, timing of migration, subsequent nest site fidelity and reproductive behaviour including egg laying with those of the remaining birds, a comparison group of 41 females. We found no effects on immediate nest survival. Fledging success and subsequent southward and northward migration patterns of the implanted birds conformed to the expectations, and arrival time on the breeding grounds in 2010–2012 did not differ from the comparison group. Compared with the comparison group, in the year after implantation, implanted birds were equally faithful to the nest site and showed equal territorial behaviour, but a paucity of behaviours indicating nests or clutches. In the 3 years after implantation, the yearly apparent survival of implanted birds was 16 % points lower. Despite intense searching, we found only three eggs of two implanted birds; all were deformed. A similarly deformed egg was reported in a similarly implanted Whimbrel Numenius phaeopus returning to breed in central Alaska. The presence in the body cavity of an object slightly smaller than a normal egg may thus lead to egg malformation and, likely, reduced egg viability. That the use of implanted satellite transmitters in these large shorebirds reduced nesting propensity and might also lead to fertility losses argues against the use of implanted transmitters for studies on breeding biology, and for a careful evaluation of the methodology in studies of migration.

Friesl

Genetic structure and viability selection in the golden eagle (Aquila chrysaetos), a vagile raptor with a Holarctic distribution

Molecular markers can reveal interesting aspects of organismal ecology and evolution, especially when surveyed in rare or elusive species. Herein, we provide a preliminary assessment of golden eagle ( Aquila chrysaetos ) population structure in North America using novel single nucleotide polymorphisms (SNPs). These SNPs included one molecular sexing marker, two mitochondrial markers, 85 putatively neutral markers that were derived from noncoding regions within large intergenic intervals, and 74 putatively nonneutral markers found in or very near protein-coding genes. We genotyped 523 eagle samples at these 162 SNPs and quantified genotyping error rates and variability at each marker. Our samples corresponded to 344 individual golden eagles as assessed by unique multilocus genotypes. Observed heterozygosity of known adults was significantly higher than of chicks, as was the number of heterozygous loci, indicating that mean zygosity measured across all 159 autosomal markers was an indicator of fitness as it is associated with eagle survival to adulthood. Finally, we used chick samples of known provenance to test for population differentiation across portions of North America and found pronounced structure among geographic sampling sites. These data indicate that cryptic genetic population structure is likely widespread in the golden eagle gene pool, and that extensive field sampling and genotyping will be required to more clearly delineate management units within North America and elsewhere.

Conservation Genetics

Lianas escape self-thinning: Experimental evidence of positive density dependence in temperate lianas Celastrus orbiculatus and C. scandens

The neighborhood density of plants strongly affects their growth, reproduction, and survival. In most cases, high density increases competition and negatively affects a focal plant in predictable ways, leading to the self-thinning law. There are, however, situations in which high densities of plants facilitate focal plant performance, resulting in positive density dependence. Despite their importance in forest gap dynamics and distinctive growth form, there have been very few studies of the effect of density on lianas or vines. We grew an invasive ( Celastrus orbiculatus ) and a native ( Celastrus scandens ) liana species together in three different density treatments, while also manipulating the light and support availability. We found that across treatment conditions, C. orbiculatus always out-performed C. scandens , showing greater relative growth rate in height and diameter, greater biomass and higher survival. Both species responded similarly to the density treatments: with plants in high density not showing a decrease in relative height growth rate compared to medium density. Aboveground biomass for C. scandens was not affected by density, while for C. orbiculatus , the most massive plants were growing in medium density without support. More surprisingly, survival analysis indicated that the two species both had significantly lower mortality rates in the highest density treatment; this trend held true across the other treatments of light and supports. More generally, this study demonstrates that these lianas can escape the consequences of high density and thus the self-thinning law that affects self-supporting plants. This suggests a broader hypothesis about lianas in general: their greater flexibility in allocating growth resources allows them to grow taller and thinner without collapsing and thereby potentially escape shading and mortality even at high densities.

Perspectives in Plant Ecology, Evolution and Syste

Emerging perspectives on resource tracking and animal movement ecology

Resource tracking, where animals increase energy gain by moving to track phenological variation in resources across space, is emerging as a fundamental attribute of animal movement ecology. However, a theoretical framework to understand when and where resource tracking should occur, and how resource tracking should lead to emergent ecological patterns, is lacking. We present a framework that unites concepts from optimal foraging theory and landscape ecology, which can be used to generate and test predictions on how resource dynamics shape animal movement across taxa, systems, and scales. Consideration of the interplay between animal movement and resource dynamics not only advances ecological understanding but can also guide biodiversity conservation in an era of global change.

Trends in Ecology & Evolution

Viewing animal migration through a social lens

Evidence of social learning is growing across the animal kingdom. Researchers have long hypothesized that social interactions play a key role in many animal migrations, but strong empirical support is scarce except in a few unique systems and species. Here, we aim to catalyze advances in the study of social migrations by synthesizing research across disciplines and providing a framework for understanding when, how, and why social influences shape the decisions animals make during migration. Integrating research across the fields of social learning and migration ecology will advance our understanding of the complex behavioral phenomena of animal migration and help to inform conservation of animal migrations in a changing world.

Trends in Ecology & Evolution

Gulf of Mexico blue hole harbors high levels of novel microbial lineages

Exploration of oxygen-depleted marine environments has consistently revealed novel microbial taxa and metabolic capabilities that expand our understanding of microbial evolution and ecology. Marine blue holes are shallow karst formations characterized by low oxygen and high organic matter content. They are logistically challenging to sample, and thus our understanding of their biogeochemistry and microbial ecology is limited. We present a metagenomic and geochemical characterization of Amberjack Hole on the Florida continental shelf (Gulf of Mexico). Dissolved oxygen became depleted at the hole’s rim (32 m water depth), remained low but detectable in an intermediate hypoxic zone (40–75 m), and then increased to a secondary peak before falling below detection in the bottom layer (80–110 m), concomitant with increases in nutrients, dissolved iron, and a series of sequentially more reduced sulfur species. Microbial communities in the bottom layer contained heretofore undocumented levels of the recently discovered phylum Woesearchaeota (up to 58% of the community), along with lineages in the bacterial Candidate Phyla Radiation (CPR). Thirty-one high-quality metagenome-assembled genomes (MAGs) showed extensive biochemical capabilities for sulfur and nitrogen cycling, as well as for resisting and respiring arsenic. One uncharacterized gene associated with a CPR lineage differentiated hypoxic from anoxic zone communities. Overall, microbial communities and geochemical profiles were stable across two sampling dates in the spring and fall of 2019. The blue hole habitat is a natural marine laboratory that provides opportunities for sampling taxa with under-characterized but potentially important roles in redox-stratified microbial processes.

Interational Society of Microbial Ecology (ISME) J

Signals of impending change

Society has an increasing awareness that there are finite limits to what we can expect the planet to absorb and still provide goods and services at current rates 1 . Both historical reconstructions and contemporary events continue to remind us that ecological regime changes are often abrupt rather than gradual. This reality motivates researchers who seek to discover leading indicators for impending ecosystem change. Berdugo et al . 2 report an important advance in our ability to anticipate the conversion of arid lands from self-organized, self-maintaining and productive ecosystems, to a state characterized by disorganization and low functionality. Such conversions have important implications for our understanding of ‘desertification’ — which is a shift from arid to desert-like conditions. Theoretical studies have suggested that patterns in the patchiness of vegetation might indicate how close a system is to making an abrupt change to desert-like conditions 3 , 4 , 5 . Empirical studies, however, have tended to show instead that simply the total cover of vegetation, rather than its arrangement, often foretells the state of the system 4 , 5 , 6 , 7 , 8 , 9 . Berdugo et al . 2 combine these competing ideas into one integrated perspective. They show how major environmental drivers, such as aridity, influence both vegetation cover and patchiness, as well as where self-organizing, stabilizing forces in the vegetation are likely to be found.

Nature Ecology & Evolution

A decade of insights into grassland ecosystem responses to global environmental change

Earth’s biodiversity and carbon uptake by plants, or primary productivity, are intricately interlinked, underlie many essential ecosystem processes, and depend on the interplay among environmental factors, many of which are being changed by human activities. While ecological theory generalizes across taxa and environments, most empirical tests of factors controlling diversity and productivity have been observational, single-site experiments, or meta-analyses, limiting our understanding of variation among site-level responses and tests of general mechanisms. A synthesis of results from ten years of a globally distributed, coordinated experiment, the Nutrient Network (NutNet), demonstrates that species diversity promotes ecosystem productivity and stability, and that nutrient supply and herbivory control diversity via changes in composition, including invasions of non-native species and extinction of native species. Distributed experimental networks are a powerful tool for tests and integration of multiple theories and for generating multivariate predictions about the effects of global changes on future ecosystems.

Nature Ecology & Evolution