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A dirty dozen ways to die: Metrics and modifiers of mortality driven by drought and warming for a tree species

Tree mortality events driven by drought and warmer temperature, often amplified by pests and pathogens, are emerging as one of the predominant climate change impacts on plants. Understanding and predicting widespread tree mortality events in the future is vital as they affect ecosystem goods and services provided by forests and woodlands, including carbon storage needed to help offset warming. Additionally, if extensive enough, tree die-off events can influence not only local climate but also climate and vegetation elsewhere via ecoclimate teleconnections. Consequently, recent efforts have focused on improving predictions of tree mortality. One of the most commercially important genera of trees is Pinus , and the most studied species globally for drought-induced tree mortality is piñon pine, Pinus edulis . Numerous metrics have been developed in association with predicting mortality thresholds or variations in mortality for this species. In this article, we compiled metrics associated with drought and warming related mortality that were developed for P. edulis or for which P. edulis was a key example species used in a calculation or prediction. We grouped these metrics into three categories: (i) those related to simple climate variables, (ii) those related to physiological responses, and (iii) those that require multi-step calculations or modeling using climate, ecohydrological, and/or ecophysiological data; and we identified the spatial-temporal scale of each of these metrics. We also compiled factors shown to modify rates or sensitivities of mortality. The metrics to predict mortality include empirical ones which often have implicit linkages to expected mechanisms, and more mechanistic ones related to physiological drivers. The metrics for P. edulis have similarities with those available for other species of Pinus . Expected future mortality events will provide an opportunity to observationally and experimentally test and compare these metrics related to tree mortality for P. edulis via near-term ecological forecasting. The metrics for P. edulis may also be useful as potential analogs for other genera. Improving predictions of tree mortality for this species and others will be increasingly important as an aid to move toward anticipatory management.

Frontiers in Forests and Global Change

Predictive framework for estimating exposure of birds to pharmaceuticals

We present and evaluate a framework for estimating concentrations of pharmaceuticals over time in wildlife feeding at wastewater treatment plants (WWTPs). The framework is composed of a series of predictive steps involving the estimation of pharmaceutical concentration in wastewater, accumulation into wildlife food items, and uptake by wildlife with subsequent distribution into, and elimination from, tissues. Because many pharmacokinetic parameters for wildlife are unavailable for the majority of drugs in use, a read-across approach was employed using either rodent or human data on absorption, distribution, metabolism, and excretion. Comparison of the different steps in the framework against experimental data for the scenario where birds are feeding on a WWTP contaminated with fluoxetine showed that estimated concentrations in wastewater treatment works were lower than measured concentrations; concentrations in food could be reasonably estimated if experimental bioaccumulation data are available; and read-across from rodent data worked better than human to bird read-across. The framework provides adequate predictions of plasma concentrations and of elimination behavior in birds but yields poor predictions of distribution in tissues. The approach holds promise, but it is important that we improve our understanding of the physiological similarities and differences between wild birds and domesticated laboratory mammals used in pharmaceutical efficacy/safety trials, so that the wealth of data available can be applied more effectively in ecological risk assessments.

Environmental Toxicology and Chemistry

Understanding the links between functional performance and environment in cold desert ecosystems through the flagship biocrust forming moss Syntrichia caninervis

Background and aims Biocrust communities shape the soil surface in drylands, regulating important ecosystem properties. Despite this, few works address key functional attributes in these poikilohydric photosynthetic communities, including the length of metabolic activity in the field and its relationship with microclimate. We aim to disentangle the links between functional performance of a keystone biocrust species and environment in drylands, in order to better understand ecosystem functioning and services provided to the soil by these organisms. Methods We developed a 23 months monitoring experiment using very high resolution measurements of chlorophyll a fluorescence and microclimate in the field, comparing habitats with soils of different particle size composition and microclimatic conditions for the flagship biocrust moss Syntrichia caninervis on the Colorado Plateau, Utah, USA. Results While growing in soils of different composition did not impair the functional performance of the species, the sun/shade plasticity did affect key physiological attributes. Our results quantified a strategy based on avoidance of the most extreme environmental conditions, where metabolic activity only occur during 19.9% of the total time monitored. Relative humidity was the best microclimatic predictor of metabolic activity, and the annual length of metabolically active periods showed strong convergences with other drylands of the world. Conclusions These results collectively suggest a strong control of the environment over the ecosystem services provided by biocrusts. Our results include evidence for an improved understanding of the plasticity of S. caninervis to its variety of habitats, and its response to potential threats under different climate change scenarios.

Utah

Exploring the pathology of an epidermal disease affecting a circum-Antarctic sea star

Over the past decade, unusual mortality outbreaks have decimated echinoderm populations over broad geographic regions, raising awareness globally of the importance of investigating such events. Echinoderms are key components of marine benthos for top-down and bottom-up regulations of plants and animals; population declines of these individuals can have significant ecosystem-wide effects. Here we describe the first case study of an outbreak affecting Antarctic echinoderms and consisting of an ulcerative epidermal disease affecting ~10% of the population of the keystone asteroid predator Odontaster validus at Deception Island, Antarctica. This event was first detected in the Austral summer 2012–2013, coinciding with unprecedented high seawater temperatures and increased seismicity. Histological analyses revealed epidermal ulceration, inflammation, and necrosis in diseased animals. Bacterial and fungal alpha diversity was consistently lower and of different composition in lesioned versus unaffected tissues (32.87% and 16.94% shared bacterial and fungal operational taxonomic units OTUs respectively). The microbiome of healthy stars was more consistent across individuals than in diseased specimens suggesting microbial dysbiosis, especially in the lesion fronts. Because these microbes were not associated with tissue damage at the microscopic level, their contribution to the development of epidermal lesions remains unclear. Our study reveals that disease events are reaching echinoderms as far as the polar regions thereby highlighting the need to develop a greater understanding of the microbiology and physiology of marine diseases and ecosystems health, especially in the era of global warming.

Scientific Reports

Higher temperature sensitivity of flowering than leaf-out alters the time between phenophases across temperate tree species

Aim The aims of this study were to evaluate the changes in the length of the time period between leaf-out and flowering across temperate tree species, and associate these changes with potential physiological and environmental drivers to enhance mechanistic insight into these phenomena. Location Central Europe. Time period 1980–2016. Major taxa studied Six temperate woody species. Methods Statistical analyses were carried out based on long-term ground observations of both spring leaf-out and flowering across temperate tree species during 1980–2016, a period characterized by rapid warming. Results The temperature sensitivity of flowering (−5.4 ± 0.04 days/℃, mean ± SE ) was higher than that of leaf-out (−4.6 ± 0.04 days/℃) across all species, regardless of whether leaf-out occurred before or after flowering. This study postulates a hypothesis attributing the different temperature sensitivities to different thermal sensitivities, thermal requirements, and photoperiodic controls. Main conclusions The larger temperature sensitivity of flowering than leaf-out resulted in an extended time period between flowering and leaf-out in species that bloom before leafing out, but a shorter time period between these phenophases in species with the opposite strategy. We would like to emphasize the importance of changes in time period between different phenophases, and we recommend conducting experimental research to reveal the underlying mechanisms of plant phenology response to climate change, and to explore its potential ecological implications.

Global Ecology and Biogeography

Mechanisms of nitrogen deposition effects on temperate forest lichens and trees

We review the mechanisms of deleterious nitrogen (N) deposition impacts on temperate forests, with a particular focus on trees and lichens. Elevated anthropogenic N deposition to forests has varied effects on individual organisms depending on characteristics both of the N inputs (form, timing, amount) and of the organisms (ecology, physiology) involved. Improved mechanistic knowledge of these effects can aid in developing robust predictions of how organisms respond to either increases or decreases in N deposition. Rising N levels affect forests in micro- and macroscopic ways from physiological responses at the cellular, tissue, and organism levels to influencing individual species and entire communities and ecosystems. A synthesis of these processes forms the basis for the overarching themes of this paper, which focuses on N effects at different levels of biological organization in temperate forests. For lichens, the mechanisms of direct effects of N are relatively well known at cellular, organismal, and community levels, though interactions of N with other stressors merit further research. For trees, effects of N deposition are better understood for N as an acidifying agent than as a nutrient; in both cases, the impacts can reflect direct effects on short time scales and indirect effects mediated through long-term soil and belowground changes. There are many gaps on fundamental N use and cycling in ecosystems, and we highlight the most critical gaps for understanding potential deleterious effects of N deposition. For lichens, these gaps include both how N affects specific metabolic pathways and how N is metabolized. For trees, these gaps include understanding the direct effects of N deposition onto forest canopies, the sensitivity of different tree species and mycorrhizal symbionts to N, the influence of soil properties, and the reversibility of N and acidification effects on plants and soils. Continued study of how these N response mechanisms interact with one another, and with other dimensions of global change, remains essential for predicting ongoing changes in lichen and tree populations across North American temperate forests.

Ecosphere

Leaf to landscape responses of giant sequoia to hotter drought: An introduction and synthesis for the special section

Hotter droughts are becoming more common as climate change progresses, and they may already have caused instances of forest dieback on all forested continents. Learning from hotter droughts, including where on the landscape forests are more or less vulnerable to these events, is critical to help resource managers proactively prepare for the future. As part of our Leaf to Landscape Project, we measured the response of giant sequoia, the world’s largest tree species, to the extreme 2012–2016 hotter drought in California. The project integrated leaf-level physiology measurements, crown-level foliage dieback surveys, and remotely sensed canopy water content (CWC) to shed light on mechanisms and spatial patterns in drought response. Here we summarize initial findings, present a conceptual model of drought response, and discuss management implications; details are presented in the other four articles of the special section on Giant Sequoias and Drought. Giant sequoias exhibited both leaf- and canopy-level responses that were effective in protecting whole-tree hydraulic integrity for the vast majority of individual sequoias. Very few giant sequoias died during the drought compared to other mixed conifer tree species; however, the magnitude of sequoia drought response varied across the landscape. This variability was partially explained by local site characteristics, including variables related to site water balance . We found that low CWC is an indicator of recent foliage dieback, which occurs when stress levels are high enough that leaf-level adjustments alone are insufficient for giant sequoias to maintain hydraulic integrity. CWC or change in CWC may be useful indicators of drought stress that reveal patterns of vulnerability to future hotter droughts. Future work will measure recovery from the drought and strengthen our ability to interpret CWC maps. Our ultimate goal is to produce giant sequoia vulnerability maps to help target management actions, such as reducing other stressors, increasing resistance to hotter drought through prescribed fire or mechanical thinning, and planting sequoias in projected future suitable habitat, which may occur outside current grove distributions. We suggest that managers compare different types of vulnerability assessments and combine vulnerability maps with other sources of information to inform decisions.

California

Increases in desert shrub productivity under elevated carbon dioxide vary with water availability

Productivity of aridland plants is predicted to increase substantially with rising atmospheric carbon dioxide (CO2) concentrations due to enhancement in plant water-use efficiency (WUE). However, to date, there are few detailed analyses of how intact desert vegetation responds to elevated CO2. From 1998 to 2001, we examined aboveground production, photosynthesis, and water relations within three species exposed to ambient (around 38 Pa) or elevated (55 Pa) CO2 concentrations at the Nevada Desert Free-Air CO2 Enrichment (FACE) Facility in southern Nevada, USA. The functional types sampled - evergreen (Larrea tridentata), drought-deciduous (Ambrosia dumosa), and winter-deciduous shrubs (Krameria erecta) - represent potentially different responses to elevated CO2 in this ecosystem. We found elevated CO2 significantly increased aboveground production in all three species during an anomalously wet year (1998), with relative production ratios (elevated:ambient CO2) ranging from 1.59 (Krameria) to 2.31 (Larrea). In three below-average rainfall years (1999-2001), growth was much reduced in all species, with only Ambrosia in 2001 having significantly higher production under elevated CO2. Integrated photosynthesis (mol CO2 m-2 y-1) in the three species was 1.26-2.03-fold higher under elevated CO2 in the wet year (1998) and 1.32-1.43-fold higher after the third year of reduced rainfall (2001). Instantaneous WUE was also higher in shrubs grown under elevated CO2. The timing of peak canopy development did not change under elevated CO2; for example, there was no observed extension of leaf longevity into the dry season in the deciduous species. Similarly, seasonal patterns in CO2 assimilation did not change, except for Larrea. Therefore, phenological and physiological patterns that characterize Mojave Desert perennials - early-season lags in canopy development behind peak photosynthetic capacity, coupled with reductions in late-season photosynthetic capacity prior to reductions in leaf area - were not significantly affected by elevated CO2. Together, these findings suggest that elevated CO2 can enhance the productivity of Mojave Desert shrubs, but this effect is most pronounced during years with abundant rainfall when soil resources are most available. ?? 2006 Springer Science+Business Media, Inc.

Ecosystems

Seasonal foraging responses of beavers to sodium-enhanced foods: An experimental assessment with field feeding trials

Salt drive is a seasonal phenomenon common to several classes of wild herbivores. Coincident with shifts of nutrient quality when plants resume growth in the spring, sodium is secondarily lost as surplus potassium is excreted. The beaver ( Castor canadensis ) is an herbivore whose dietary niche closely follows that of other herbivores that are subject to salt drive, but no published studies to date have assessed the likelihood of its occurrence. To quantify if beavers experience seasonal salt drive, we designed a field experiment to measure the foraging responses of beavers to sodium-enhanced foods. We used sodium-treated (salted) and control (no salt) food items (aspen [ Populus tremuloides ] and pine [ Pinus spp.] sticks) during monthly feeding trials at beaver-occupied wetlands. If conventional ontogeny of salt drive was operant, we expected to observe greater utility of sodium-treated food items by beavers in May and June. Further, if water lilies ( Nymphaea spp. and Nuphar spp.) supply beavers with sodium to meet dietary requirements as is widely speculated, we expected foraging responses to sodium-treated food items at wetlands where water lilies were absent to be greater than at wetlands where water lily was present. Aspen was selected by beavers in significantly greater amounts than pine. There was no difference between the mean percent consumed of salted and control aspen sticks by beavers at lily and non-lily wetlands, and no differences in temporal consumption associated with salted or control pine sticks at either wetland type. Salted pine was consumed in greater amounts than unsalted pine. We propose that the gastrointestinal or renal physiology of beavers may preclude solute loss, thereby preventing salt drive.

Journal of Mammalogy

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

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

Alaska

Assessment of treated wastewater infiltration in Bright Angel Wash and the potential for contaminants of emerging concern influencing spring water quality along the South Rim of the Grand Canyon in Grand Canyon National Park, Arizona

In April 2021, a synoptic study conducted by the U.S. Geological Survey (USGS) and National Park Service (NPS) identified wastewater-related contaminants of emerging concern (CECs) in springs along the South Rim of the Grand Canyon. These springs are located north of Bright Angel Wash, an ephemeral channel that receives treated effluent from the South Rim Wastewater Treatment Plant (SRWTP). Although water flows southwest and away from the canyon, there is evidence that treated wastewater is finding a flow path along fractures associated with the Bright Angel Fault back to water sources along the South Rim. The CECs identified during the April 2021 sampling included several per- and polyfluoroalkyl substances (PFAS) and pharmaceutical compounds. The PFAS compounds detected only consisted of perfluoroalkyl acids, and these were only detected at Bright Angel Wash (treated wastewater), Monument Spring, and upper Horn Bedrock Spring. The other five sampled springs (the Salt Creek, Horn East Alluvium, Garden, Pumphouse, and Pipe Springs) had no detections of PFAS compounds. The five perfluoroalkyl acids detected at Monument Spring (in descending order of concentration) were perfluorobutanesulfonic acid (PFBS), perfluoropentanoic acid, perfluorooctanoic acid, perfluorohexanoic acid, and perfluorooctanesulfonic acid. Only the short-chained PFBS and perfluorobutanoic acid compounds were detected at the upper Horn Bedrock Spring. All the same perfluoroalkyl acids were found at Bright Angel Wash, except for PFBS. Monument Spring was the only spring to have detections of pharmaceuticals. The two pharmaceuticals detected at the highest concentrations at Monument Spring were diphenhydramine (antihistamine) and carbamazepine (anticonvulsant or anti-epileptic drug). The other detected pharmaceuticals included (in descending order of concentration) fluconazole (antifungal), sulfamethoxazole (antibiotic), metformin (diabetes drug), tramadol (opioid analgesic), and venlafaxine (antidepressant and nerve pain medication). The same pharmaceuticals were detected in the wastewater at Bright Angel Wash but in greater concentrations. No CEC concentrations measured at Monument Spring exceeded any drinking water standards or human health benchmarks; however, most of the compounds detected have no regulatory standards. Studies of the ecological effects of these compounds show that some of the compounds detected can have endocrine and physiological effects, but generally, effects were observed at concentrations multiple orders of magnitude greater than what was measured during the April 2021 study. Data from 1980 through 2022 retrieved from the Water Quality Portal were combined with data from the one-time synoptic sampling in April 2021 to assess the usefulness of other analytes for identifying a wastewater connection to South Rim springs. Most of the historical water chemistry data showed a statistical difference between samples collected within and east of the Garden Creek watershed and samples collected from locations in watersheds to the west of Garden Creek, including the Horn Creek, Salt Creek, Monument Creek, and Hermit Creek watersheds, which roughly align with the Bright Angel Fault. Most of the historical analytes were inconclusive as potential wastewater tracers, but nitrate, chloride, and gadolinium data possibly support the historical contribution of wastewater to Monument Spring.

Arizona

Near-field receiving-water monitoring of trace metals and a benthic community near the Palo Alto Regional Water Quality Control Plant in south San Francisco Bay, California—2020

Trace-metal concentrations in sediment and in the clam Limecola petalum (World Register of Marine Species, 2020; formerly reported as Macoma balthica and M. petalum ), clam reproductive activity, and benthic macroinvertebrate community structure were investigated in a mudflat 1 kilometer (km) south of the discharge of the Palo Alto Regional Water Quality Control Plant (PARWQCP) in south San Francisco Bay, California. This report includes the data collected by the U.S. Geological Survey (USGS) for January 2020–December 2020 (Cain and others, 2022). These data append to long-term datasets extending back to 1974. A major focus of the report is an integrated description of the 2020 data within the context of the longer, multidecadal dataset. This dataset supports the City of Palo Alto’s Near-Field Receiving- Water Monitoring Program, initiated in 1994. Silver and copper contamination substantially decreased at the site in the 1980s following the implementation by PARWQCP of advanced wastewater-treatment and source-control measures. Since the 1990s, concentrations of these elements in surface sediments have continued to decrease, although more slowly. For example, from 1994 to 2020, the minimum annual mean silver concentration—0.20 milligram per kilogram (mg/kg)—was observed in multiple years. In 2020, silver concentrations ranged from 0.18 to 0.28 mg/kg. These concentrations are 2 to 3 times higher than the regional background concentration. Presently (2020), sediment-copper concentrations appear to be near the regional background level. Over the same period (1994–2020), sedimentary iron and zinc exhibited modest decreases. Sedimentary aluminum, chromium, mercury, nickel, and selenium have not exhibited any trend. Since 1994, silver and copper concentrations in L. petalum have varied seasonally, apparently in response to a combination of site-specific metal exposures and cyclic growth and reproduction, as reported previously. Seasonal patterns for other elements, including chromium, mercury, nickel, selenium, and zinc, generally were similar in timing and magnitude as those for silver and copper. Downward trends in the silver and zinc concentrations in L. petalum during 1994–2020 were evident and appeared to be related to the general physiological condition of the clam, indicated by a condition index. Biological effects of elevated silver and copper contamination at the Palo Alto site have been interpreted from data collected during and after the recession of these contaminants. Concentrations of both elements in the soft tissues of L. petalum decreased with sedimentary copper and silver. This pattern was associated with changes in the reproductive activity of L. petalum , as well as the structure of the benthic invertebrate community. Reproductive activity of L. petalum increased as metal concentrations in L. petalum decreased (Hornberger and others, 2000), and presently is stable with almost all animals initiating reproduction in the fall and spawning the following spring. Analyses of the benthic community structure indicate that the infaunal invertebrate community has shifted from one dominated by several opportunistic species when silver and copper exposures were highest to one in which the species abundance is more evenly distributed, a pattern that indicates a more stable community that is subjected to fewer stressors. Importantly, this long-term change is unrelated to other metals and other measured environmental factors, including salinity and sediment composition. In addition, two of the opportunistic species ( Ampelisca abdita and Streblospio benedicti ) that brood their young and live on the surface of the sediment in tubes have shown a continual decrease in dominance coincident with the decrease in metals. Both species had short-lived rebounds in abundance in 2008, 2009, and 2010 and showed signs of increasing abundance in 2020. Heteromastus filiformis (a subsurface polychaete worm that lives in the sediment, consumes sediment and organic particles residing in the sediment, and reproduces by laying its eggs on or in the sediment) showed a concurrent increase in dominance and, in the last several years before 2008, showed a stable population. H. filiformis abundance increased slightly from 2011 to 2012 and returned to pre-2011 numbers in 2020. The reproductive mode of most species that were present in 2020 was indicative of species that were capable of movement either as pelagic larvae or as mobile adults. Although oviparous species were lower in number in this group, the authors hypothesize that these species will return slowly as more species move back into the area. The use of functional ecology was highlighted in the 2020 benthic community data, which showed that the animals that have now returned to the mudflat are those that can respond successfully to a physical, nontoxic disturbance. Today, community data show a mix of species that consume the sediment, or filter feed, those that have pelagic larvae that must survive landing on the sediment, and those that brood their young. The long-term recovery observed after the 1970s can be ascribed to the decrease in sediment pollutants.

California

Leaf gas exchange characteristics of three neotropical mangrove species in response to varying hydroperiod

We determined how different hydroperiods affected leaf gas exchange characteristics of greenhouse-grown seedlings (2002) and saplings (2003) of the mangrove species Avicennia germinans (L.) Stearn., Laguncularia racemosa (L.) Gaertn. f., and Rhizophora mangle L. Hydroperiod treatments included no flooding (unflooded), intermittent flooding (intermittent), and permanent flooding (flooded). Plants in the intermittent treatment were measured under both flooded and drained states and compared separately. In the greenhouse study, plants of all species maintained different leaf areas in the contrasting hydroperiods during both years. Assimilation–light response curves indicated that the different hydroperiods had little effect on leaf gas exchange characteristics in either seedlings or saplings. However, short-term intermittent flooding for between 6 and 22 days caused a 20% reduction in maximum leaf-level carbon assimilation rate, a 51% lower light requirement to attain 50% of maximum assimilation, and a 38% higher demand from dark respiration. Although interspecific differences were evident for nearly all measured parameters in both years, there was little consistency in ranking of the interspecific responses. Species by hydroperiod interactions were significant only for sapling leaf area. In a field study, R. mangle saplings along the Shark River in the Everglades National Park either demonstrated no significant effect or slight enhancement of carbon assimilation and water-use efficiency while flooded. We obtained little evidence that contrasting hydroperiods affect leaf gas exchange characteristics of mangrove seedlings or saplings over long time intervals; however, intermittent flooding may cause short-term depressions in leaf gas exchange. The resilience of mangrove systems to flooding, as demonstrated in the permanently flooded treatments, will likely promote photosynthetic and morphological adjustment to slight hydroperiod shifts in many settings. .

Tree Physiology

Monitoring responses to variation in food supply for a migratory waterfowl: American Black Duck (Anas rubripes) in winter

Wintering Black Ducks ( Anas rubripes ) concentrate in wetlands along the Atlantic coast where natural and anthropogenic disturbances have increased over the last 50 years, a period in which the population of Black Ducks has declined. We studied the sensitivity of Black Ducks to perturbations in food supply that often result from disturbances by storms, predators, and people. In the paper, we characterize the responses of captive Black Ducks to shifts in food quality and availability during winter and apply those measures to a comparison of wild birds. Captive ducks that were fed intermittently (3 consecutive days/week) compensated for fasted days to achieve similar body mass and body fat to control birds that were fed every day on both animal- and plant-based diets. However, birds that were fed intermittently expended 15% more energy each day than controls when both groups were fed (536 vs. 464 kJ/kg 0.75 ), which indicates that variable food supply increases the costs of maintenance and thus reduces the number of birds that can be supported on the same resource of food without interruptions to foraging. Egg production was not affected by diet quality provided in spring or by the frequency of feeding during the preceding winter months. Black Ducks lost body fat through winter in captivity and in the wild. Fat stores of birds in New Jersey were greater than those of birds in Maine (13.3 vs. 8.3% of body mass) in January, which reflected the high energy demands of cold temperatures in Maine. Values for ∂ 15 N were greater in Maine than in New Jersey for both red blood cells and plasma, which indicated a consistent diet of marine invertebrates in Maine. Greater isotopic variation in red blood cells indicated that diets were more diverse in New Jersey than in Maine for both ∂ 15 N (9.7 ± 1.1 vs. 11.2 ± 0.4‰) and for ∂ 13 C (− 15.1 ± 2.2 vs. − 13.8 ± 1.4‰). Plasma ∂ 13 C was enriched over red blood cells in wild birds especially those with low fat stores, which suggested birds with low energy stores were shifting diets. Black Ducks can compensate for disturbances in feeding by increasing intakes if they have access to high quality wetlands where they are able to find abundant food. High energy demands at cold temperatures may constrain fat stores and thus the tolerance of feeding disturbances especially at the northern limits of the winter range. We hypothesize that decreasing variation in diet may indicate an increase in vulnerability to disturbance in winter when body fat is low. Recent efforts to assess and improve habitat quality of Black Ducks could be enhanced by monitoring the body composition and diet of birds to assess their vulnerability to disturbances in food supply and energy demands.

Journal of Comparative Physiology B: Biochemical,

Early-warning signals of individual tree mortality based on annual radial growth

Tree mortality is a key driver of forest dynamics and its occurrence is projected to increase in the future due to climate change. Despite recent advances in our understanding of the physiological mechanisms leading to death, we still lack robust indicators of mortality risk that could be applied at the individual tree scale. Here, we build on a previous contribution exploring the differences in growth level between trees that died and survived a given mortality event to assess whether changes in temporal autocorrelation, variance, and synchrony in time-series of annual radial growth data can be used as early warning signals of mortality risk. Taking advantage of a unique global ring-width database of 3065 dead trees and 4389 living trees growing together at 198 sites (belonging to 36 gymnosperm and angiosperm species), we analyzed temporal changes in autocorrelation, variance, and synchrony before tree death (diachronic analysis), and also compared these metrics between trees that died and trees that survived a given mortality event (synchronic analysis). Changes in autocorrelation were a poor indicator of mortality risk. However, we found a gradual increase in inter-annual growth variability and a decrease in growth synchrony in the last ∼20 years before mortality of gymnosperms, irrespective of the cause of mortality. These changes could be associated with drought-induced alterations in carbon economy and allocation patterns. In angiosperms, we did not find any consistent changes in any metric. Such lack of any signal might be explained by the relatively high capacity of angiosperms to recover after a stress-induced growth decline. Our analysis provides a robust method for estimating early-warning signals of tree mortality based on annual growth data. In addition to the frequently reported decrease in growth rates, an increase in inter-annual growth variability and a decrease in growth synchrony may be powerful predictors of gymnosperm mortality risk, but not necessarily so for angiosperms.

Frontiers in Plant Science