Global change effects on Bromus tectorum L. (Poaceae) at its high‐elevation range margin

Global change is likely to affect invasive species distribution, especially at range margins. In the eastern Sierra Nevada, California, USA, the invasive annual grass, Bromus tectorum, is patchily distributed and its impacts have been minimal compared with other areas of the Intermountain West. We used a series of in situ field manipulations to determine how B. tectorum might respond to changing climatic conditions and increased nitrogen deposition at the high‐elevation edge of its invaded range. Over 3 years, we used snow fences to simulate changes in snowpack, irrigation to simulate increased frequency and magnitude of springtime precipitation, and added nitrogen (N) at three levels (0, 5, and 10 g m^?2) to natural patches of B. tectorum growing under the two dominant shrubs, Artemisia tridentata and Purshia tridentata, and in intershrub spaces (INTR). We found that B. tectorum seedling density in April was lower following deeper snowpack possibly due to delayed emergence, yet there was no change in spikelet production or biomass accumulation at the time of harvest. Additional spring rain events increased B. tectorum biomass and spikelet production in INTR plots only. Plants were primarily limited by water in 2009, but colimited by N and water in 2011, possibly due to differences in antecedent moisture conditions at the time of treatments. The threshold at which N had an effect varied with magnitude of water additions. Frequency of rain events was more influential than magnitude in driving B. tectorum growth and fecundity responses. Our results suggest that predicted shifts from snow to rain could facilitate expansion of B. tectorum at high elevation depending on timing of rain events and level of N deposition. We found evidence for P‐limitation at this site and an increase in P‐availability with N additions, suggesting that stoichiometric relationships may also influence B. tectorum spread.     

Snowpack,fire, and forest disturbance: interactions affect montane invasions by non‐native shrubs

Montane regions worldwide have experienced relatively low plant invasion rates, a trend attributed to increased climatic severity, low rates of disturbance, and reduced propagule pressure relative to lowlands. Manipulative experiments at elevations above the invasive range of non‐native species can clarify the relative contributions of these mechanisms to montane invasion resistance, yet such experiments are rare. Furthermore, global climate change and land use changes are expected to cause decreases in snowpack and increases in disturbance by fire and forest thinning in montane forests. We examined the importance of these factors in limiting montane invasions using a field transplant experiment above the invasive range of two non‐native lowland shrubs, Scotch broom (Cytisus scoparius) and Spanish broom (Spartium junceum), in the rain–snow transition zone of the Sierra Nevada of California. We tested the effects of canopy closure, prescribed fire, and winter snow depth on demographic transitions of each species. Establishment of both species was most likely at intermediate levels of canopy disturbance, but at this intermediate canopy level, snow depth had negative effects on winter survival of seedlings. We used matrix population models to show that an 86% reduction in winter snowfall would cause a 2.8‐fold increase in population growth rates in Scotch broom and a 3.5‐fold increase in Spanish broom. Fall prescribed fire increased germination rates, but decreased overall population growth rates by reducing plant survival. However, at longer fire return intervals, population recovery between fires is likely to keep growth rates high, especially under low snowpack conditions. Many treatment combinations had positive growth rates despite being above the current invasive range, indicating that propagule pressure, disturbance, and climate can all strongly affect plant invasions in montane regions. We conclude that projected reductions in winter snowpack and increases in forest disturbance are likely to increase the risk of invasion from lower elevations.     

Eco‐evolutionary responses of Bromus tectorum to climate change: implications for biological invasions

How plant populations, communities, and ecosystems respond to climate change is a critical focus in ecology today. The responses of introduced species may be especially rapid. Current models that incorporate temperature and precipitation suggest that future Bromus tectorum invasion risk is low for the Colorado Plateau. With a field warming experiment at two sites in southeastern Utah, we tested this prediction over 4 years, measuring B. tectorum phenology, biomass, and reproduction. In a complimentary greenhouse study, we assessed whether changes in field B. tectorum biomass and reproductive output influence offspring performance. We found that following a wet winter and early spring, the timing of spring growth initiation, flowering, and summer senescence all advanced in warmed plots at both field sites and the shift in phenology was progressively larger with greater warming. Earlier green‐up and development was associated with increases in B. tectorum biomass and reproductive output, likely due early spring growth, when soil moisture was not limiting, and a lengthened growing season. Seeds collected from plants grown in warmed plots had higher biomass and germination rates and lower mortality than seeds from ambient plots. However, in the following two dry years, we observed no differences in phenology between warmed and ambient plots. In addition, warming had a generally negative effect on B. tectorum biomass and reproduction in dry years and this negative effect was significant in the plots that received the highest warming treatment. In contrast to models that predict negative responses of B. tectorum to warmer climate on the Colorado Plateau, the effects of warming were more nuanced, relied on background climate, and differed between the two field sites. Our results highlight the importance of considering the interacting effects of temperature, precipitation, and site‐specific characteristics such as soil texture, on plant demography and have direct implications for B. tectorum invasion dynamics on the Colorado Plateau.     

Soil water exploitation after fire: competition between Bromus tectorum (cheatgrass) and two native species

Summary Causes for the widespread abundance of the alien grass Bromus tectorum (cheatgrass) after fire in semiarid areas of western North America may include: (1) utilization of resources freed by the removal of fireintolerant plants; and (2) successful competition between B. tectorum and individual plants that survive fire. On a site in northwestern Nevada (USA), measurements of soil water content, plant water potential, aboveground biomass production, water use efficiency, and B. tectorum tiller density were used to determine if B. tectorum competes with either of two native species (Stipa comata and Chrysothamnus viscidiflorus) or simply uses unclaimed resources. Soil water content around native species occurring with B. tectorum was significantly lower (P<0.05) than around individuals without B. tectorum nearby. Native species had significantly more negative plant water potential when they occurred with B. tectorum. Aboveground biomass was significantly higher for native species without B. tectorum. However, the carbon isotope ratio of leaves for native species with B. tectorum was not significantly different from individuals without B. tectorum. Thus, B. tectorum competes with native species for soil water and negatively affects their wate status and productivity, but the competition for water does not affect water use efficiency of the native species. These adverse effects of B. tectorum competition on the productivity and water status of native species are also evident at 12 years after a fire. This competitive ability of B. tectorum greatly enhances its capability to exploit soil resources after fire and to enhance its status in the community.     

Evaluating Sierra Nevada Yellow-Legged Frog Distribution Using Environmental DNA

Genetic tools that identify species from trace DNA samples could supplement traditional survey methods to clarify distributional limits of rare species. For species with legal habitat protection, elevational limits of distributions are used to determine where management actions may affect endangered species. The endangered Sierra Nevada yellow-legged frog (Rana sierrae) generally is found down to 1,370 m, but in the Plumas National Forest, California, USA, there are a number of historical records below this elevation, resulting in protections extending to 1,067 m. This species is phenotypically similar to the foothill yellow-legged frog (R. boylii), with which it occasionally hybridizes. We used a combination of genetic methods to investigate the fine-scale distribution of the Sierra Nevada yellow-legged frog in the Plumas National Forest. We collected and analyzed environmental DNA (eDNA) samples from all accessible lower elevation sites with records of Sierra Nevada yellow-legged frog (n = 17) and swabbed 220 individuals for genetic identification from 2016–2018 to clarify the distribution of this endangered species. We created a climatic suitability model using the validated Sierra Nevada yellow-legged frog records and current (1970–2000) climate models to assess additional highly suitable localities for Sierra Nevada yellow-legged frog presence using eDNA capture. We did not confirm detection of Sierra Nevada yellow-legged frog eDNA at any historical sites and identified all swabbed individuals from below 1,370 m (n = 144) as foothill yellow-legged frogs. We located a new Sierra Nevada yellow-legged frog site (at 1,919 m) during surveys guided by the climatic suitability model. It does not appear after extensive eDNA and genetic sampling that the Sierra Nevada yellow-legged frog occurs below 1,370 m in this portion of their range at present. Our results show that eDNA sampling can be used as an effective management tool to evaluate historical locations and previously unknown suitable localities for current presence of a species of interest. © 2021 The Authors. The Journal of Wildlife Management published by Wiley Periodicals LLC on behalf of The Wildlife Society.     

OBSERVATIONS ON SNOW ALGAE IN CALIFORNIA1,2

Algae that impart a red color to snowfields are rather common in California. Red snow occurs mainly in the Sierra Nevada at altitudes of 10,000–12,000 ft (3050–3600 in) and can occur at high altitudes where snow persists in other parts of the state. The distribution in the Sierra was similar in 1969 and 1970, contrasting snowfall years. Colored snow was found from May to October in old, wet snow-fields. The predominant color was red and occurred as surface patches in depressions in the snow. The color could extend as deep as 30 cm below the snow surface. Algae in the snowfields of the Tioga Pass area (Sierra Nevada) were large, red, spherical cells of Chlamydomonas nivalis. No other algae were seen. Their distribution, as measured by cell numbers and chlorophyll a, was patchy. Algal cells and chlorophyll a were mainly distributed at or near the snow surface but extended down to a depth of 10 cm. Light intensity was greatly attenuated by snow, but enough light for photosynthesis was found at 50 cm below the surface. Nutrient content of one snow sample was very low. The populations were very actively photosynthetic and took up as much as 65% of added ¹⁴CO₂ in only 3 hr. It was tentatively concluded that CO₂ limits in situ photosynthesis. Photosynthesis was inhibited by melting snow samples. Rough calculations of the growth rate suggested in situ generation times of only a few days for these algae.     

Direct Effects of Soil Amendments on Field Emergence and Growth of the Invasive Annual Grass Bromus tectorum L. and the Native Perennial Grass Hilaria jamesii (Torr.) Benth

Bromus tectorum L. is a non-native, annual grass that has invaded western North America. In SE Utah, B. tectorum generally occurs in grasslands dominated by the native perennial grass, Hilaria jamesii (Torr.) Benth. and rarely where the natives Stipa hymenoides Roem. and Schult. and S. comata Trin. & Rupr. are dominant. This patchy invasion is likely due to differences in soil chemistry. Previous laboratory experiments investigated using soil amendments that would allow B. tectorum to germinate but would reduce B. tectorum emergence without affecting H. jamesii. For this study we selected the most successful treatments (CaCl₂, MgCl₂, NaCl and zeolite) from a previous laboratory study and applied them in the field in two different years at B. tectorum-dominated field sites. All amendments except the lowest level of CaCl₂ and zeolite negatively affected B. tectorum emergence and/or biomass. No amendments negatively affected the biomass of H. jamesii but NaCl reduced emergence. Amendment effectiveness depended on year of application and the length of time since application. The medium concentration of zeolite had the strongest negative effect on B. tectorum with little effect on H. jamesii. We conducted a laboratory experiment to determine why zeolite was effective and found it released large amounts of Na⁺, adsorbed Ca²⁺, and increased Zn²⁺, Fe²⁺, Mn²⁺, Cu²⁺, exchangeable Mg²⁺, exchangeable K, and NH₄⁺ in the soil. Our results suggest several possible amendments to control B. tectorum. However, variability in effectiveness due to abiotic factors such as precipitation and soil type must be accounted for when establishing management plans.     

The ecology of mosquito larvae (Diptera: Culicidae) in the subalpine zone of the eastern Sierra Nevada Mountains,California, U.S.A.

Mosquito larvae were collected from the subalpine region of the eastern Sierra Nevada Mountains from 2011 to 2014. Two watersheds were sampled and sites selected were mainly vernal snow‐melt pools and wet meadows. Seven Aedes species, Culiseta incidens (Thomson), and Culex tarsalis Coquillett were collected. The most abundant and widely distributed species were Ae. hexodontus Dyar and Ae. tahoensis Dyar. Aedes tahoensis was the predominate species in woodland snow‐melt habitats. Some species were found at most elevations while others were found more often at specific elevations. The most restrictive species was Ae. ventrovittis Dyar which occurred almost exclusively between 3,219 m a.s.l. and 3,390 m a.s.l. Shannon and Simpson species diversity indices demonstrated that species diversity was greater in meadow habitats compared to woodland habitats. Mixed woodland/meadows, rock pools, and shallow grass pools were intermediate in species diversity. Abiotic factors such as snowpack and water temperature impacted species development times and when habitats dried. It was concluded that spatial and temporal patterns of habitats, along with elevation, influenced species presence and larval development. The results of the present study and previous work in the eastern Sierras will help guide future research that focuses on the potential change in the distribution and seasonality of subalpine mosquitoes and disease potential in the eastern Sierras as climatic conditions change.     

Seedling establishment of two shrubby plants native to the Sierra Nevada mountain range

The Sierra Nevada mountain range near the Mediterranean Sea is an unique environment known for the variety of endemic species. Nevertheless, an alpine ski station situated on the mountain has dramatically affected the landscape, leaving some areas barren. In an effort to restore the vegetation cover, laboratory experiments were conducted with seeds of Genista versicolor Boiss and Reseda complicata Bory, two shrubby plants native to Sierra Nevada. Using different concentrations of two plant growth regulators, Ethrel and N⁶-benzyladenine, seeds from both species were planted in soil sampled from the alpine ski resort. Surprisingly, both Ethrel and N⁶-benzyladenine significantly improved seedling establishment. Consequently, seedling pre-treatment with definite plant growth regulators could be a useful approach to revegetation of the Sierra Nevada mountain range.     

Variation in phenotypic plasticity for native and invasive populations of Bromus tectorum

Phenotypic plasticity is often considered important for invasive plant success, yet relatively few studies have assessed plasticity in both native and invasive populations of the same species. We examined the plastic response to temperature for Bromus tectorum populations collected from similar shrub-steppe environments in the Republics of Armenia and Georgia, where it is native, and along an invasive front in California and Nevada. Plants were grown in growth chambers in either ‘warm’ (30/20 °C, day/night) or ‘cold’ (10/5 °C) conditions. Invasive populations exhibited greater adaptive plasticity than natives for freezing tolerance (as measured by chlorophyll a fluorescence), such that invasive populations grown in the cold treatment exhibited the highest tolerance. Invasive populations also exhibited more rapid seedling emergence in response to warm temperatures compared to native populations. The climatic conditions of population source locations were related to emergence timing for invasive populations and to freezing tolerance across all populations combined. Plasticity in growth-related traits such as biomass, allocation, leaf length, and photosynthesis did not differ between native and invasive populations. Rather, some growth-related traits were very plastic across all populations, which may help to dampen differences in biomass in contrasting environments. Thus, invasive populations were found to be particularly plastic for some important traits such as seedling emergence and freezing tolerance, but plasticity at the species level may also be an important factor in the invasive ability of B. tectorum.     

Population structure and genetic diversity of Bromus tectorum within the small grain production region of the Pacific Northwest

Bromus tectorum L. is an invasive winter annual grass naturalized across the United States. Numerous studies have investigated B. tectorum population structure and genetics in the context of B. tectorum as an ecological invader of natural areas and rangeland. Despite the wealth of information regarding B. tectorum, previous studies have not focused on, or made comparisons to, B. tectorum as it persists in individual agroecosystems. The objectives of this study were to assess the genetic diversity and structure, the occurrence of generalist and specialist genotypes, and the influence of climate on distribution of B. tectorum sourced exclusively from within small grain production regions of the Pacific Northwest. Genetic diversity of B. tectorum sourced from agronomic fields was found to be similar to what has been observed from other land use histories. Six distinct genetic clusters of B. tectorum were identified, with no evidence to indicate that any of the genetic clusters were better adapted to a particular geographical area or climate within the region. Given the apparent random spatial distribution of B. tectorum genetic clusters at the spatial scale of this analysis, unique genotypes may be well mixed within region, similar to what was reported for other inbreeding weedy grass species.     

Vascular plant diversity and climate change in the upper zone of Sierra Nevada,Spain

Abstract

This study examines the effects of altitudinal, temperature and aspect gradients on vascular plant species richness on mountain tops in Sierra Nevada (Spain) at different spatial scales (1 m² quadrats, plot clusters of 4 m², upper summit area down to the 5-m contour line, entire summit down to the 10-m contour line). The methodology follows the Global Observation Research Initiative in Alpine Environments (GLORIA) programme. Floristic and soil temperature data of eight summits sites in two neighbouring regions of the high part of Sierra Nevada (from 2668 m to 3327 m a.s.l.) were used in this study. In total, 102 taxa were recorded (84 genera; 29 families). The species richness decreased, whereas the proportion of endemic taxa increased with elevation. There were significant linear relationships between species richness and altitude and average soil temperature at each spatial scale. However, there was no significant relationship between species richness and aspect variables. Facing continued climate change, the high-altitude flora of Sierra Nevada is expected to be particularly vulnerable and prone to warming-induced biodiversity losses due to the high proportion of endemic taxa, ranging from 23% at lower elevations up to 67% at higher ones.     

Impact of early root competition on fitness components of four semiarid species

Summary Plant demographic and root exclusion approaches were used to examine the influence of roots of adult Artemisia tridentata, Agropyron desertorum, and Agropyron spicatum individuals on seedling survival of four C₃ semiarid species, three perennials, Ar. tridentata, Ag. desertorum, Ag. spicatum, and an annual, Bromus tectorum. Furthermore, height of Ar. tridentata seedlings and seed production of B. tectorum were assessed. The probability of a seedling being alive significantly depended on the seedling species, the neighboring adult species, and on the depth to which root competition was excluded. As seedlings, survival of Agropyron species did not differ, whereas survival of Ar. tridentata seedlings was higher than Ag. desertorum and was similar to Ag. spicatum. Bromus tectorum maintained significantly higher survival rates than perennial seedlings. Established individuals of Ar. tridentata reduced seedling survival more than established individuals of either Agropyron species. Seedling survival significantly increased with greater depth of root exclusion for the perennials but did not significantly affect seedling survival of B. tectorum. Height of Ar. tridentata seedlings and seed production of B. tectorum significantly increased with depth of root exclusion. Seed production of B. tectorum was highest when competing with Ag. desertorum and was lowest with Ar. tridentata. Root competition decreased the seed population of B. tectorum in the next generation even though it had no impact on survival. Competition in the upper soil horizon occurs between seedlings and established adults early in the growing season and potentially restricts root growth of seedlings. In arid and semiarid ecosystems, soil moisture is depleted from the upper horizons first, resulting in the death of seedlings that do not have access to moisture.     

Contrasting climate influences on Nothofagus pumilio establishment along elevational gradients

Predicted warmer temperatures and more frequent extreme climatic events in the southern Andes may affect the dynamics of the Patagonian forests. These environmental changes may differentially alter the probability of Nothofagus pumilio establishment across its altitudinal range of distribution. We monitored fruit fall, seedling emergence and survival at altitudinal distribution range of N. pumilio forests in Santa Cruz (49°22′ S—72°56′ W), Argentina. Fruit fall, seedling emergence and survival were tested in relation to drought, based on the SPEI (Standardized Precipitation Evapotranspiration Index), interacting with elevation. Fruit fall was consistently higher at low elevation and the abundance of fruits was not affected by drought intensity. Density of new-born seedlings was?~?9 to 24 times lower at low- than at high-elevation plots in our first observations (2014 and 2016), characterized by warm-dry climatic conditions in spring-early summers. In contrast, seedling abundance was?~?1.5 times larger at low elevations during relatively cold-wet growing seasons. Survival probability was explained by the interaction between SPEI and elevation. At low elevation and in dry periods, survival probability was lower (CI 54–72%) than in wet periods (CI 68–84%) but at high elevation similar survival was registered even with positive or negative SPEI values. Our results show interacting effects of elevation and drought on tree establishment at the elevation limits, with positive and negative drought effects at high and low elevations, respectively. Predicted increase of extreme drought events during the XXI century could be detrimental for N. pumilio establishment at dry, low-elevation forests.

     

Climate change disrupts local adaptation and favours upslope migration

Contemporary climate change is proceeding at an unprecedented rate. The question remains whether populations adapted to historical conditions can persist under rapid environmental change. We tested whether climate change will disrupt local adaptation and reduce population growth rates using the perennial plant Boechera stricta (Brassicaceae). In a large‐scale field experiment conducted over five years, we exposed > 106 000 transplants to historical, current, or future climates and quantified fitness components. Low‐elevation populations outperformed local populations under simulated climate change (snow removal) across all five experimental gardens. Local maladaptation also emerged in control treatments, but it was less pronounced than under snow removal. We recovered local adaptation under snow addition treatments, which reflect historical conditions. Our results revealed that low elevation populations risk rapid decline, whereas upslope migration could enable population persistence and expansion at higher elevation locales. Local adaptation to historical conditions could increase vulnerability to climate change, even for geographically widespread species.     

Climate structures genetic variation across a species' elevation range: a test of range limits hypotheses

Gene flow may influence the formation of species range limits, and yet little is known about the patterns of gene flow with respect to environmental gradients or proximity to range limits. With rapid environmental change, it is especially important to understand patterns of gene flow to inform conservation efforts. Here we investigate the species range of the selfing, annual plant, Mimulus laciniatus, in the California Sierra Nevada. We assessed genetic variation, gene flow, and population abundance across the entire elevation‐based climate range. Contrary to expectations, within‐population plant density increased towards both climate limits. Mean genetic diversity of edge populations was equivalent to central populations; however, all edge populations exhibited less genetic diversity than neighbouring interior populations. Genetic differentiation was fairly consistent and moderate among all populations, and no directional signals of contemporary gene flow were detected between central and peripheral elevations. Elevation‐driven gene flow (isolation by environment), but not isolation by distance, was found across the species range. These findings were the same towards high‐ and low‐elevation range limits and were inconsistent with two common centre‐edge hypotheses invoked for the formation of species range limits: (i) decreasing habitat quality and population size; (ii) swamping gene flow from large, central populations. This pattern demonstrates that climate, but not centre‐edge dynamics, is an important range‐wide factor structuring M. laciniatus populations. To our knowledge, this is the first empirical study to relate environmental patterns of gene flow to range limits hypotheses. Similar investigations across a wide variety of taxa and life histories are needed.     

Effects of the duration of cold stratification on early life stages of the Mediterranean alpine plant Silene ciliata

Cold stratification provided by snow cover is essential to break seed dormancy in many alpine plant species. The forecast reduction in snow precipitation and snow cover duration in most temperate mountains as a result of global warming could threaten alpine plant populations, especially those at the edge of their species distribution, by altering the dynamics of early life stages. We simulated some effects of a reduction in the snow cover period by manipulating the duration of cold stratification in seeds of Silene ciliata, a Mediterranean alpine specialist. Seeds from three populations distributed along an altitudinal gradient were exposed to different periods of cold stratification (2, 4 and 6 months) in the laboratory and then moved to common garden conditions in a greenhouse. The duration of the cold stratification treatment and population origin significantly affected seed emergence percentage, emergence rate and seedling size, but not the number of seedling leaves. The 6‐month and 4‐month cold stratification treatments produced higher emergence percentages and faster emergence rates than seeds without cold stratification treatment. No significant cold stratification duration x seed population origin interactions were found, thus differential sensitivity to cold stratification along elevation is not supported.     

Past and future demographic dynamics of alpine species: limited genetic consequences despite dramatic range contraction in a plant from the Spanish Sierra Nevada

Anthropogenic global climate change is expected to cause severe range contractions among alpine plants. Alpine areas in the Mediterranean region are of special concern because of the high abundance of endemic species with narrow ranges. This study combined species distribution models, population structure analyses and Bayesian skyline plots to trace the past and future distribution and diversity of Linaria glacialis, an endangered narrow endemic species that inhabits summits of Sierra Nevada (Spain). The results showed that: (i) the habitat of this alpine‐Mediterranean species in Sierra Nevada suffered little changes during glacial and interglacial stages of late Quaternary; (ii) climatic oscillations in the last millennium (Medieval Warm Period and Little Ice Age) moderately affected the demographic trends of L. glacialis; (iii) future warming conditions will cause severe range contractions; and (iv) genetic diversity will not diminish at the same pace as the distribution range. As a consequence of the low population structure of this species, genetic impoverishment in the alpine zones of Sierra Nevada should be limited during range contraction. We conclude that maintenance of large effective population sizes via high mutation rates and high levels of gene flow may promote the resilience of alpine plant species when confronted with global warming.     

Genetic evidence for the persistence of the critically endangered Sierra Nevada red fox in California

California is home to both the native state-threatened Sierra Nevada red fox (Vulpes vulpes necator), which historically inhabited high elevations of the Sierra Nevada and Cascade mountains, and to multiple low-elevation red fox populations thought to be of exotic origin. During the past few decades the lowland populations have dramatically expanded their distribution, and possibly moved into the historic range of the native high-elevation fox. To determine whether the native red fox persists in its historic range in California, we compared mitochondrial cytochrome-b haplotypes of the only currently-known high-elevation population (n = 9 individuals) to samples from 3 modern lowland populations (n = 35) and historic (1911–1941) high-elevation (n = 22) and lowland (n = 7) populations. We found no significant population differentiation among the modern and historic high-elevation populations (average pairwise F _ST = 0.06), but these populations differed substantially from all modern and historic lowland populations (average pairwise F _ST = 0.52). Among lowland populations, the historic and modern Sacramento Valley populations were not significantly differentiated from one another (F _ST = −0.06), but differed significantly from recently founded populations in the San Francisco Bay region and in southern California (average pairwise F _ST = 0.42). Analysis of molecular variance indicated that 3 population groupings (mountain, Sacramento Valley, and other lowland regions) explained 45% of molecular variance (F _CT = 0.45) whereas only 4.5% of the variance was partitioned among populations within these groupings (F _SC = 0.08). These findings provide strong evidence that the native Sierra Nevada red fox has persisted in northern California. However, all nine samples from this population had the same haplotype, suggesting that several historic haplotypes may have become lost. Unidentified barriers have apparently prevented gene flow from the Sacramento Valley population to other eastern or southern populations in California. Future studies involving nuclear markers are needed to assess the origin of the Sierra Nevada red fox and to quantify levels of nuclear gene flow.     

Best environmental predictors of breeding phenology differ with elevation in a common woodland bird species

Temperatures in mountain areas are increasing at a higher rate than the Northern Hemisphere land average, but how fauna may respond, in particular in terms of phenology, remains poorly understood. The aim of this study was to assess how elevation could modify the relationships between climate variability (air temperature and snow melt‐out date), the timing of plant phenology and egg‐laying date of the coal tit (Periparus ater). We collected 9 years (2011–2019) of data on egg‐laying date, spring air temperature, snow melt‐out date, and larch budburst date at two elevations (~1,300 m and ~1,900 m asl) on a slope located in the Mont‐Blanc Massif in the French Alps. We found that at low elevation, larch budburst date had a direct influence on egg‐laying date, while at high‐altitude snow melt‐out date was the limiting factor. At both elevations, air temperature had a similar effect on egg‐laying date, but was a poorer predictor than larch budburst or snowmelt date. Our results shed light on proximate drivers of breeding phenology responses to interannual climate variability in mountain areas and suggest that factors directly influencing species phenology vary at different elevations. Predicting the future responses of species in a climate change context will require testing the transferability of models and accounting for nonstationary relationships between environmental predictors and the timing of phenological events.