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Robust age estimation of southern sea otters from multiple morphometrics

Reliable age estimation is an essential tool to assess the status of wildlife populations and inform successful management. Aging methods, however, are often limited by too few data, skewed demographic representation, and by single or uncertain morphometric relationships. In this study, we synthesize age estimates in southern sea otters Enhydra lutris nereis from 761 individuals across 34 years of study, using multiple noninvasive techniques and capturing all life stages from 0 to 17 years of age. From wild, stranded, and captive individuals, we describe tooth eruptions, tooth wear, body length, nose scarring, and pelage coloration across ontogeny and fit sex‐based growth functions to the data. Dental eruption schedules provided reliable and identifiable metrics spanning 0.3–9 months. Tooth wear was the most reliable predictor of age of individuals aged 1–15 years, which when combined with total length, explained >93% of observed age. Beyond age estimation, dental attrition also indicated the maximum lifespan of adult teeth is 13‒17 years, corresponding with previous estimates of life expectancy. Von Bertalanffy growth function model simulations of length at age gave consistent estimates of asymptotic lengths (male L oo = 126.0‒126.8 cm, female L oo = 115.3‒115.7 cm), biologically realistic gestation periods ( t 0 = 115 days, SD = 10.2), and somatic growth (male k = 1.8, SD = 0.1; female k = 2.1, SD = 0.1). Though exploratory, we describe how field radiographic imaging of epiphyseal plate development or fusions may improve aging of immature sea otters. Together, our results highlight the value of integrating information from multiple and diverse datasets to help resolve conservation problems.

Ecology and Evolution

Assessment of spatial genetic structure to identify populations at risk for infection of an emerging epizootic disease

Understanding the geographic extent and connectivity of wildlife populations can provide important insights into the management of disease outbreaks but defining patterns of population structure is difficult for widely distributed species. Landscape genetic analyses are powerful methods for identifying cryptic structure and movement patterns that may be associated with spatial epizootic patterns in such cases. We characterized patterns of population substructure and connectivity using microsatellite genotypes from 2,222 white-tailed deer ( Odocoileus virginianus ) in the Mid-Atlantic region of the United States, a region where chronic wasting disease was first detected in 2009. The goal of this study was to evaluate the juxtaposition between population structure, landscape features that influence gene flow, and current disease management units. Clustering analyses identified four to five subpopulations in this region, the edges of which corresponded to ecophysiographic provinces. Subpopulations were further partitioned into 11 clusters with subtle ( F ST ≤ 0.041), but significant genetic differentiation. Genetic differentiation was lower and migration rates were higher among neighboring genetic clusters, indicating an underlying genetic cline. Genetic discontinuities were associated with topographic barriers, however. Resistance surface modeling indicated that gene flow was diffuse in homogenous landscapes, but the direction and extent of gene flow were influenced by forest cover, traffic volume, and elevational relief in subregions heterogeneous for these landscape features. Chronic wasting disease primarily occurred among genetic clusters within a single subpopulation and along corridors of high landscape connectivity. These results may suggest a possible correlation between population substructure, landscape connectivity, and the occurrence of diseases for widespread species. Considering these factors may be useful in delineating effective management units, although only the largest features produced appreciable differences in subpopulation structure. Disease mitigation strategies implemented at the scale of ecophysiographic provinces are likely to be more effective than those implemented at finer scales.

Maryland, Pennsylvania, Virginia

Supervised versus unsupervised approaches to classification of accelerometry data

Sophisticated animal-borne sensor systems are increasingly providing novel insight into how animals behave and move. Despite their widespread use in ecology, the diversity and expanding quality and quantity of data they produce have created a need for robust analytical methods for biological interpretation. Machine learning tools are often used to meet this need. However, their relative effectiveness is not well known and, in the case of unsupervised tools, given that they do not use validation data, their accuracy can be difficult to assess. We evaluated the effectiveness of supervised ( n = 6), semi-supervised ( n = 1), and unsupervised ( n = 2) approaches to analyzing accelerometry data collected from critically endangered California condors ( Gymnogyps californianus ). Unsupervised K-means and EM (expectation–maximization) clustering approaches performed poorly, with adequate classification accuracies of <0.8 but very low values for kappa statistics (range: −0.02 to 0.06). The semi-supervised nearest mean classifier was moderately effective at classification, with an overall classification accuracy of 0.61 but effective classification only of two of the four behavioral classes. Supervised random forest (RF) and k-nearest neighbor (kNN) machine learning models were most effective at classification across all behavior types, with overall accuracies >0.81. Kappa statistics were also highest for RF and kNN, in most cases substantially greater than for other modeling approaches. Unsupervised modeling, which is commonly used for the classification of a priori-defined behaviors in telemetry data, can provide useful information but likely is instead better suited to post hoc definition of generalized behavioral states. This work also shows the potential for substantial variation in classification accuracy among different machine learning approaches and among different metrics of accuracy. As such, when analyzing biotelemetry data, best practices appear to call for the evaluation of several machine learning techniques and several measures of accuracy for each dataset under consideration.

Ecology and Evolution

Improving the accessibility and transferability of machine learning algorithms for identification of animals in camera trap images: MLWIC2

Motion‐activated wildlife cameras (or “camera traps”) are frequently used to remotely and noninvasively observe animals. The vast number of images collected from camera trap projects has prompted some biologists to employ machine learning algorithms to automatically recognize species in these images, or at least filter‐out images that do not contain animals. These approaches are often limited by model transferability, as a model trained to recognize species from one location might not work as well for the same species in different locations. Furthermore, these methods often require advanced computational skills, making them inaccessible to many biologists. We used 3 million camera trap images from 18 studies in 10 states across the United States of America to train two deep neural networks, one that recognizes 58 species, the “species model,” and one that determines if an image is empty or if it contains an animal, the “empty‐animal model.” Our species model and empty‐animal model had accuracies of 96.8% and 97.3%, respectively. Furthermore, the models performed well on some out‐of‐sample datasets, as the species model had 91% accuracy on species from Canada (accuracy range 36%–91% across all out‐of‐sample datasets) and the empty‐animal model achieved an accuracy of 91%–94% on out‐of‐sample datasets from different continents. Our software addresses some of the limitations of using machine learning to classify images from camera traps. By including many species from several locations, our species model is potentially applicable to many camera trap studies in North America. We also found that our empty‐animal model can facilitate removal of images without animals globally. We provide the trained models in an R package (MLWIC2: Machine Learning for Wildlife Image Classification in R), which contains Shiny Applications that allow scientists with minimal programming experience to use trained models and train new models in six neural network architectures with varying depths.

Ecology and Evolution

Development of a genotyping-in-thousands by sequencing (GT-seq) panel for identifying individuals and estimating relatedness among Alaska black bears (Ursus americanus)

The management and conservation of large mammals, such as black bears ( Ursus americanus ), have long been informed by genetic estimates of population size and individual dispersal. Amplicon sequencing methods, also known as ‘genotyping-in-thousands-by sequencing’ (GT-seq), now enable the efficient and cost-effective genotyping of hundreds of loci and individuals in the same sequencing run. Here, we develop a GT-seq panel for individual identification and kinship inference in Alaska black bears. Using genomic data from restriction site-associated DNA sequencing of hunter-harvested bears from Southcentral Alaska ( n = 85), we identified 170 microhaplotype and single nucleotide polymorphism (SNP) loci that were highly heterozygous in local populations. To enable sexing of individuals, we also included a previously published sex-linked locus in the GT-seq panel. We empirically validated the GT-seq panel using samples collected at different spatial scales. These samples included tissues ( n = 82) obtained from bears within a small geographic area in Anchorage, Alaska, which were likely to be relatives as well as the hunter-harvested samples collected from geographically widespread locations throughout Southcentral Alaska. Empirical validation indicated high genotyping success and genotype reproducibility across replicate subsamples. Computer simulations demonstrated that the GT-seq panel had ample statistical power for distinguishing distinct individuals and first-order relatives (parent-offspring and full-sibling pairs) from unrelated individuals. As a final proof of concept, the panel was used to identify individual bears and close kin sampled from urban and wild habitats in Anchorage, Alaska. We anticipate that the GT-seq panel will be a useful genomic resource for the monitoring and management of Alaska black bear populations. ons.

Alaska

Citizen science data collection for integrated wildlife population analyses

Citizen science, or community science, has emerged as a cost-efficient method to collect data for wildlife monitoring. To inform research and conservation, citizen science sampling designs should collect data that match the robust statistical analyses needed to quantify species and population patterns. Further increasing the contributions of citizen science, integrating citizen science data with other datasets and datatypes can improve population estimates and expand the spatiotemporal extent of inference. We demonstrate these points with a citizen science program called iSeeMammals developed in New York state in 2017 to supplement costly systematic spatial capture-recapture sampling by collecting opportunistic data from one-off observations, hikes, and camera traps. iSeeMammals has initially focused on the growing population of American black bear ( Ursus americanus ), with integrated analysis of iSeeMammals camera trap data with systematic data for a region with a growing bear population. The triumvirate of increased spatial and temporal coverage by at least twofold compared to systematic sampling, an 83% reduction in annual sampling costs, and improved density estimates when integrated with systematic data highlight the benefits of collecting presence-absence data in citizen science programs for estimating population patterns. Additional opportunities will come from applying presence-only data, which are oftentimes more prevalent than presence-absence data, to integrated models. Patterns in data submission and filtering also emphasize the importance of iteratively evaluating patterns in engagement, usability, and accessibility, especially focusing on younger adult and teenage demographics, to improve data quality and quantity. We explore how the development and use of integrated models may be paired with citizen science project design in order to facilitate repeated use of datasets in standalone and integrated analyses for supporting wildlife monitoring and informing conservation.

Frontiers in Ecology and Evolution

Backpack satellite transmitters reduce survival but not nesting propensity or success of greater sage-grouse

Telemetry technology is ubiquitous for studying the behavior and demography of wildlife, including the use of traditional very high frequency (VHF) radio telemetry and more recent methods that record animal locations using global positioning systems (GPS). Satellite-based GPS telemetry allows researchers to collect high spatial–temporal resolution data remotely but may also come with additional costs. For example, recent studies from the southern Great Basin suggested GPS transmitters attached via backpacks may reduce the survival of greater sage-grouse ( Centrocercus urophasianus ) relative to VHF transmitters attached via collars that have been in use for decades. While some evidence suggests GPS backpacks reduce survival, no studies have examined the effects of GPS backpacks on breeding behavior and success. Therefore, we compared survival, breeding behavior, and nest success of sage-grouse hens marked with both VHF collars and GPS backpack transmitter over a 7-year period in central Idaho, USA. GPS backpacks reduced spring–summer survival of sage-grouse hens relative to hens with VHF collars, where daily mortality probability was 68%–82% higher from March 1 to August 1. Yet satellite GPS backpacks did not consistently affect nest success or the likelihood or timing of nest initiation relative to VHF collars. Daily nest survival varied annually and with timing of nest initiation and nest age, but marginal effects of transmitter type were statistically insignificant and interactions between transmitter type and study year produced no meaningful patterns. Our results corroborate recent studies for the effect of satellite GPS backpacks on sage-grouse survival, but also suggest that these transmitters do not appear to affect components of fecundity. Our results therefore add important context to recent debate surrounding the effects of GPS backpacks on sage-grouse, and the relative strengths and weaknesses of different transmitter types for understanding behavior and population dynamics.

Idaho

Guidelines for a priori grouping of species in hierarchical community models

Recent methodological advances permit the estimation of species richness and occurrences for rare species by linking species-level occurrence models at the community level. The value of such methods is underscored by the ability to examine the influence of landscape heterogeneity on species assemblages at large spatial scales. A salient advantage of community-level approaches is that parameter estimates for data-poor species are more precise as the estimation process borrows from data-rich species. However, this analytical benefit raises a question about the degree to which inferences are dependent on the implicit assumption of relatedness among species. Here, we assess the sensitivity of community/group-level metrics, and individual-level species inferences given various classification schemes for grouping species assemblages using multispecies occurrence models. We explore the implications of these groupings on parameter estimates for avian communities in two ecosystems: tropical forests in Puerto Rico and temperate forests in northeastern United States. We report on the classification performance and extent of variability in occurrence probabilities and species richness estimates that can be observed depending on the classification scheme used. We found estimates of species richness to be most precise and to have the best predictive performance when all of the data were grouped at a single community level. Community/group-level parameters appear to be heavily influenced by the grouping criteria, but were not driven strictly by total number of detections for species. We found different grouping schemes can provide an opportunity to identify unique assemblage responses that would not have been found if all of the species were analyzed together. We suggest three guidelines: (1) classification schemes should be determined based on study objectives; (2) model selection should be used to quantitatively compare different classification approaches; and (3) sensitivity of results to different classification approaches should be assessed. These guidelines should help researchers apply hierarchical community models in the most effective manner.

Ecology and Evolution

Coastal vegetation responses to large dam removal on the Elwha River

Introduction: Large dam removals provide a restoration opportunity for shrinking coastal wetland habitats. Dam removal can increase sediment delivery to sediment-starved river deltas and estuaries by restoring natural sediment transport and mobilizing reservoir-impounded sediment. However, rapid mobilization of massive quantities of sediment stored behind large dams also constitutes a major ecological perturbation. Information is lacking on coastal habitat responses to sediment pulses of this magnitude. Methods: Removal of two large dams along the Elwha River (Washington, USA) in 2011–2014 released ~20.5 Mt of impounded sediment, ~5.4 Mt of which were deposited in the delta and estuary (hereafter, delta). We used time series of aerial imagery, digital elevation models, and vegetation field sampling to examine plant community responses to this sediment pulse across seven years during and after dam removal. Results: Between 2011 and 2018, the Elwha River delta increased by ~26.8 ha. Vegetation colonized ~16.4 ha of new surfaces, with mixed pioneer vegetation on supratidal beach, river bars, and river mouth bars and emergent marsh vegetation in intertidal aquatic habitats. Colonization occurred on surfaces that were higher and more stable in elevation and farther from the shoreline. Compared to established delta plant communities, vegetation on new surfaces had lower cover of dominant species and functional groups, with very low woody cover, and lower graminoid cover than dunegrass and emergent marsh communities. Over time following surface stabilization, however, vegetation on new surfaces increased in species richness, cover, and similarity to established communities. By 2018, ~1.0 ha of vegetation on new surfaces had developed into dunegrass or willow–alder communities and ~5.9 ha had developed into emergent marsh. At the same time, dam removal had few discernible effects on established delta plant communities. Discussion: Together, these results suggest that rapid sediment mobilization during large dam removal has potential to expand coastal wetland habitat without negatively affecting established plant communities. However, as sediment loads declined in 2016–2018, new delta surfaces decreased by ~4.5 ha, and ~1.6 ha of new vegetation reverted to no vegetation. Long-term persistence of the expanded coastal habitat will depend on ongoing erosional and depositional processes under the restored natural sediment regime.

Washington

Estimation bias in water-quality constituent concentrations and fluxes: A synthesis for Chesapeake Bay rivers and streams

Flux quantification for riverine water-quality constituents has been an active area of research. Statistical approaches are often employed to make estimation for days without observations. One such approach is the Weighted Regressions on Time, Discharge, and Season (WRTDS) method. While WRTDS has been used in many investigations, there is a general lack of effort to identify factors that influence its estimation bias. This work was aimed to (1) synthesize and compare WRTDS estimation bias for constituent concentrations and fluxes for rivers and streams in the Chesapeake Bay watershed (including headwater sites) and (2) identify controlling factors from five broad categories (watershed size, sampling practice, concentration and discharge conditions, land use, and geology). Five major constituents were considered, namely, suspended sediment (SS), total phosphorus (TP), total nitrogen (TN), orthophosphate (PO 4 ), and nitrate-plus-nitrite (NO x ). For both concentration and flux, estimation bias follows the general order of SS > TP > PO 4 > TN ≈ NO x . Median TN and NO x bias statistics were near zero, with an equal distribution of small positive and negative bias. TP, PO 4 , and SS each showed a median positive bias across sites of <18% for flux and <7% for concentration. Particulate constituents, especially SS, tend to have larger bias at sites with smaller sampling frequencies, shorter sampling record lengths, and smaller watershed sizes. Results of multivariate models showed that both flux and concentration biases are most affected by concentration and discharge variabilities and the length of concentration record. In comparison, flux bias of particulate constituents is more affected by flow variability, whereas flux bias of dissolved constituents is more affected by concentration variability. Moreover, analysis using classification and regression trees provided additional information on how the factors affected flux bias: when all site-constituent combinations are considered, large flux biases are more likely associated with sites that have large concentration and discharge variabilities, small lengths of concentration record, and small sampling frequencies. These results may be useful for identifying sites with large biases, modifying monitoring practice at existing sites to reduce those biases, and choosing new monitoring locations in the Chesapeake watershed and beyond.

Chesapeake Bay watershed

Southern California Bight 2003 Regional Monitoring Program: V. water quality

More than $30 million is expended annually on environmental monitoring in the Southern California Bight (SCB), yet only 5% of the Bight is monitored on an ongoing basis. Therefore, environmental managers in the SCB decided to expand their monitoring program and, starting in 1994, decided to conduct periodic regional assessments of ecosystem condition and assess the overall health of the SCB. Sixty-five different organizations collaborated in 2003 to create the third SCB Regional Monitoring Program (Bight '03). Bight '03 was designed to be integrated regional monitoring program that encompasses regulatory, academic, and non-governmental agencies. Bight '03 had three components: Coastal Ecology, Shoreline Microbiology, and Water Quality. This report addresses the purpose, approach, findings, and recommendations from the Water Quality component, which focused on contamination-laden stormwater runoff, in particularly its variability in time and space as well as its short-term ecological impacts. Specifically, the Bight '03 Water Quality component had three primary goals, the first of which was to described the temporal evolution of stormwater plumes produced by the major southern California rivers. Specifically, the study was intended to determine how far offshore the plumes extended, how rapidly they advected, how long before the plumes dispersed and how these properties differed among storms and river systems. The second goal was to describe how the physical properties (e.g., turbidity, temperature, salinity) of the plume related to biogeochemical and ecological properties that are of more direct concern to the water quality management community. Accomplished primarily through ship-based sampling of water quality parameters, this second goal was to describe how far offshore, and for how ;long after the storm, elevated bacterial concentrations, toxicity, and nutrients could be detected. Similar to the fist goal, the study also addressed how these answers differed among storms and river systems. The final goal was to determine whether relationships between environmental indicators derived from coincident satellite remote sensing and in situ data sets are sufficiently robust for remote sensing to become a routine water quality monitoring programs. Remote sensing data potentially provide coastal managers with synoptic near-real time regional information about prevailing ocean conditions and hazards that would complement existing field-based sampling protocols, but only if there is a thorough understanding of how to interpret and utilize the proxy measures, such as ocean color. The understanding of these priorities through Bight '03 sampling is intended to provide the basis for developing more efficient, widespread and coast-effective coastal ocean monitoring techniques. Water quality data were collected across eight major river systems within four geographic regions of southern California. Field measurements included the primary contaminants of interest, i.e., bacterial concentrations, water toxicity, and nutrients, as well as related parameters such as temperatures, salinity, total suspended solids, transmissivity, chlorophyll, and colored dissolved organic material (CDOM) concentrations. For each of the four major regions, i.e., Santa Clara/Ventura Rivers, Ballona Creek/Santa Monica Bay, San Pedro Shelf, and the San Diego, Tijuana Rivers, two stormwater events were sampled for up to three days by ship resulting in 574 water column CTD+ profiles and 705 discrete water samples during 36 ship-days. These data were analyzed in combination with MODIS ocean color satellite remote sensing, buoy meteorological observations, drifters, and HF radar current measurements to evaluate the dispersal patterns, dynamics, and impacts of the freshwater runoff plumes. Based on these data and resulting analyses, the principal conclusions were as follow: - Stormwater runoff turbidity plumes were found to be spatially extensive, covering up to 2500 km 2 within the Southern California Bight nearshore zone, and persisting over the entire duration of the post-storm sampling period (at least 3 days). - The spatial and temporal extent of the portion of the plume with contaminants was far less than that of the turbidity plume, typically representing <10% of its area (30-70% off Tijuana); however, with contaminant impacts generally greatly reduced or absent by the third or fourth day of sampling - Pseudo-nitzschia , a harmful algae that produces domoic acid, was found to be more abundant than previously reported. - Accurately describing stormwater runoff plumes requires a combination of in situ and remote sensing assessment tools, with satellite data providing valuable synoptic information. From these conclusions, the following recommendations are provided: - Future studies designed to describe stormwater plumes should include a combination of ship - and remote sensing-based methods. - CDOM is a good proxy of the freshwater runoff plume and should be added as a standard measurement parameter on water quality instrument packages. - Investigations are needed that assess on a local basis the spatial extent of ecological effects of stormwater plumes early in the storm, ideally accompanied by airborne imagery to provide improved temporal & spatial resolution, to fill in knowledge gaps. The next Bight regional monitoring program should focus on quantifying nutrient loadings and dynamics in association with stormwater runoff and other sources, and characterize their attendant ecosystem impacts such as phytoplankton blooms.

California