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Analysis and Management of Animal Populations: Modeling, Estimation and Decision Making

This book deals with the processes involved in making informed decisions about the management of animal populations. It covers the modeling of population responses to management actions, the estimation of quantities needed in the modeling effort, and the application of these estimates and models to the development of sound management decisions. The book synthesizes and integrates in a single volume the methods associated with these themes, as they apply to ecological assessment and conservation of animal populations. KEY FEATURES * Integrates population modeling, parameter estimation and * decision-theoretic approaches to management in a single, cohesive framework * Provides authoritative, state-of-the-art descriptions of quantitative * approaches to modeling, estimation and decision-making * Emphasizes the role of mathematical modeling in the conduct of science * and management * Utilizes a unifying biological context, consistent mathematical notation, * and numerous biological examples

Book

Integrated species distribution model using historical data shows decline in a common semi-aquatic mammal

Effective conservation requires an understanding of drivers of a species' distribution as well as long-term changes in their distribution. In recent decades, advances in data collection and analysis have allowed researchers to integrate a wide range of information to model species distributions, particularly by allowing presence-only data and detection-nondetection data to be formally combined in integrated species distribution models (ISDMs). However, these models are rarely used to investigate long-term trends, which are important in evaluating a species' status. Here, we use historical presence-only data of river otters ( Lontra canadensis ; 366 latrine locations from 1999 to 2007 and 105 locations of road-killed individuals recorded from 1999 to 2020) and 919 detection-nondetection surveys from 230 sites between 2021 and 2023 to understand the current distribution of river otters in Rhode Island, USA, as well as the changes in river otter distribution over the past two decades. We found that river otters were strongly associated with key habitat features such as streams and water, positively associated with urban areas, and tolerant of some contaminants, such as lead. Furthermore, despite uncertainties in historical river otter occurrence, we found clear supporting evidence that river otter intensity of use had declined from 1999 to 2023. This decline occurred despite being protected from harvest and in contrast to range expansions in other parts of the northeastern USA throughout the second half of the 20th century. Our results suggest the utility of this approach to detect declines in species for which historical data are available and a need for better understanding the cause of river otter declines. Where monitoring consists of opportunistically collected data, species conservation could benefit by continuing to collect these data as well as introducing designed surveys, as this would allow better integration of data types, improving trend estimation and reducing the amount of (typically more expensive) designed surveys needed.

Rhode Island

The dominance of introduced plant species in the diets of migratory Galapagos tortoises increases with elevation on a human-occupied island

The distribution of resources and food selection are fundamental to the ecology, life history, physiology, population dynamics, and conservation of animals. Introduced plants are changing foraging dynamics of herbivores in many ecosystems often with unknown consequences. Galapagos tortoises, like many herbivores, undertake migrations along elevation gradients driven by variability in vegetation productivity which take them into upland areas dominated by introduced plants. We sought to characterize diet composition of two species of Galapagos tortoises, focussing on how the role of introduced forage species changes over space and the implications for tortoise conservation. We quantified the distribution of tortoises with elevation using GPS telemetry. Along the elevation gradient, we quantified the abundance of introduced and native plant species, estimated diet composition by recording foods consumed by tortoises, and assessed tortoise physical condition from body weights and blood parameter values. Tortoises ranged between 0 and 429 m in elevation over which they consumed at least 64 plant species from 26 families, 44 percent of which were introduced species. Cover of introduced species and the proportion of introduced species in tortoise diets increased with elevation. Introduced species were positively selected for by tortoises at all elevations. Tortoise physical condition was either consistent or increased with elevation at the least biologically productive season on Galapagos. Santa Cruz tortoises are generalist herbivores that have adapted their feeding behavior to consume many introduced plant species that has likely made a positive contribution to tortoise nutrition. Some transformed habitats that contain an abundance of introduced forage species are compatible with tortoise conservation.

Galapagos Islands

Beyond overlap: Considering habitat preference and fitness outcomes in the umbrella species concept

Umbrella species and other surrogate species approaches to conservation provide an appealing framework to extend the reach of conservation efforts beyond single species. For the umbrella species concept to be effective, populations of multiple species of concern must persist in areas protected on behalf of the umbrella species. Most assessments of the concept, however, focus exclusively on geographic overlap among umbrella and background species, and not measures that affect population persistence (e.g. habitat quality or fitness). We quantified the congruence between the habitat preferences and nesting success of a high-profile umbrella species (greater sage-grouse, Centrocercus urophasianus , hereafter ‘sage-grouse’), and three sympatric species of declining songbirds (Brewer's sparrow Spizella breweri , sage thrasher Oreoscoptes montanus and vesper sparrow Pooecetes gramineus ) in central Wyoming, USA during 2012–2013. We used machine-learning methods to create data-driven predictions of sage-grouse nest-site selection and nest survival probabilities by modeling field-collected sage-grouse data relative to habitat attributes. We then used field-collected songbird data to assess whether high-quality sites for songbirds aligned with those of sage-grouse. Nest sites selected by songbirds did not coincide with sage-grouse nesting preferences, with the exception that Brewer's sparrows preferred similar nest sites to sage-grouse in 2012. Moreover, the areas that produced higher rates of songbird nest survival were unrelated to those for sage-grouse. Our findings suggest that management actions at local scales that prioritize sage-grouse nesting habitat will not necessarily enhance the reproductive success of sagebrush-associated songbirds. Measures implemented to conserve sage-grouse and other purported umbrella species at broad spatial scales likely overlap the distribution of many species, however, broad-scale overlap may not translate to fine-scale conservation benefit beyond the umbrella species itself. The maintenance of microhabitat heterogeneity important for a diversity of species of concern will be critical for a more holistic application of the umbrella species concept.

Animal Conservation

Future directions to escalate benefits of stepping-stone approach for conservation translocations

Through a reintroduction case study on the critically endangered Vancouver Island marmot (Marmota vancouverensis), we introduced a ‘stepping-stone’ approach which utilizes the transition of released individuals among populations to maximize demographic growth potential (Lloyd et al. 2019). We greatly appreciate and hereby reflect on the thoughtful commentaries by Chauvenet (2019), Hayward (2019) and Thévenin (2019) to propose 3 key topics for future exploration.

Animal Conservation

Estimating golden-cheeked warbler immigration: Implications for the spatial scale of conservation

Understanding the factors that drive population dynamics is fundamental to species conservation and management. Since the golden-cheeked warbler S etophaga chrysoparia was first listed as endangered, much effort has taken place to monitor warbler abundance, occupancy, reproduction and survival. Yet, despite being directly related to local population dynamics, movement rates have not been estimated for the species. We used an integrated population model to investigate the relationship between immigration rate, fledging rate, survival probabilities and population growth rate for warblers in central Texas, USA. Furthermore, using a deterministic projection model, we examined the response required by vital rates to maintain a viable population across varying levels of immigration. Warbler abundance fluctuated with an overall positive trend across years. In the absence of immigration, the abundance would have decreased. However, the population could remain viable without immigration if both adult and juvenile survival increased by almost half or if juvenile survival more than doubled. We also investigated the response required by fledging rates across a range of immigration in order to maintain a viable population. Overall, we found that immigration was required to maintain warbler target populations, indicating that warbler conservation and management programs need to be implemented at larger spatial scales than current efforts to be effective. This study also demonstrates that by using limited data within integrated population models, biologists are able to monitor multiple key demographic parameters simultaneously to gauge the efficacy of strategies designed to maximize warbler viability in a changing landscape.

Animal Conservation

Presence-nonpresence surveys of golden-cheeked warblers: detection, occupancy and survey effort

Surveys to detect the presence or absence of endangered species may not consistently cover an area, account for imperfect detection or consider that detection and species presence at sample units may change within a survey season. We evaluated a detection?nondetection survey method for the federally endangered golden-cheeked warbler (GCWA) Dendroica chrysoparia. Three study areas were selected across the breeding range of GCWA in central Texas. Within each area, 28-36 detection stations were placed 200 m apart. Each detection station was surveyed nine times during the breeding season in 2 consecutive years. Surveyors remained up to 8 min at each detection station recording GCWA detected by sight or sound. To assess the potential influence of environmental covariates (e.g. slope, aspect, canopy cover, study area) on detection and occupancy and possible changes in occupancy and detection probabilities within breeding seasons, 30 models were analyzed. Using information-theoretic model selection procedures, we found that detection probabilities and occupancy varied among study areas and within breeding seasons. Detection probabilities ranged from 0.20 to 0.80 and occupancy ranged from 0.56 to 0.95. Because study areas with high detection probabilities had high occupancy, a conservative survey effort (erred towards too much surveying) was estimated using the lowest detection probability. We determined that nine surveys of 35 stations were needed to have estimates of occupancy with coefficients of variation <20%. Our survey evaluation evidently captured the key environmental variable that influenced bird detection (GCWA density) and accommodated the changes in GCWA distribution throughout the breeding season.

Animal Conservation

Migration Mapper: Identifying movement corridors and seasonal ranges for large mammal conservation

Modern tracking technology has facilitated a novel understanding of terrestrial mammal movement while revealing that movements are being truncated and lost. The first step towards conserving mobile animals is identifying movement corridors and key seasonal ranges. Yet, the identification and subsequent mapping of these important areas has remained a challenge due to the analytical skills necessary to conduct such analyses. Migration Mapper (MM) is a user-friendly software that provides tools to analyse global positioning system (GPS) collar data to create season-specific, population-level polygons representing areas where most of a population moves (i.e. movement corridors) and areas where most of a population spends time (e.g. high-use areas, seasonal ranges). MM consists of six standalone modules including data cleaning and review, seasonal movement delineation, movement model application, calculation of population-level outputs and visualization of results. Analysis of GPS data using MM can provide the spatial polygons necessary to facilitate conservation and policy planning. New initiatives at the local and global levels are already beginning to use MM to facilitate conservation of large, terrestrial mammals.

Methods in Ecology and Evolution

Monitoring landscape-level distribution and migration Phenology of Raptors using a volunteer camera-trap network

Conservation of animal migratory movements is among the most important issues in wildlife management. To address this need for landscape-scale monitoring of raptor populations, we developed a novel, baited photographic observation network termed the “Appalachian Eagle Monitoring Program” (AEMP). During winter months of 2008–2012, we partnered with professional and citizen scientists in 11 states in the United States to collect approximately 2.5 million images. To our knowledge, this represents the largest such camera-trap effort to date. Analyses of data collected in 2011 and 2012 revealed complex, often species-specific, spatial and temporal patterns in winter raptor movement behavior as well as spatial and temporal resource partitioning between raptor species. Although programmatic advances in data analysis and involvement are needed, the continued growth of the program has the potential to provide a long-term, cost-effective, range-wide monitoring tool for avian and terrestrial scavengers during the winter season. Perhaps most importantly, by relying heavily on citizen scientists, AEMP has the potential to improve long-term interest and support for raptor conservation and serve as a model for raptor conservation programs in other portions of the world.

Wildlife Society Bulletin

Decision analysis for habitat conservation of an endangered, range-limited salamander

Many species of conservation concern are habitat limited and often a major focus of management for these species is habitat acquisition and/or restoration. Deciding the location of habitat restoration or acquisition to best benefit a protected species can be a complicated subject with competing management objectives, ecological uncertainties and stochasticity. Structured decision making (SDM) could be a useful approach for explicitly incorporating those complexities while still working toward species conservation and/or recovery. We applied an SDM approach to Red Hills salamander Phaeognathus hubrichti habitat conservation decision making. Phaeognathus hubrichti is a severely range-limited endemic species in south central Alabama and has highly specific habitat requirements. Many known populations live on private lands and the primary mode of habitat protection is habitat conservation planning, but such plans are non-binding and not permanent. Working with stakeholders, we developed an objectives hierarchy linking land acquisition or protection actions to fundamental objectives. We built a model to assess and compare the quality of the habitat in the known range of P. hubrichti . Our model evaluated key habitat attributes of 5814 pixels of 1 km 2 each and ranked the pixels from best to worst with respect to P. hubrichti habitat requirements. Our results are a spatially explicit valuation of each pixel, with respect to its probable benefit to P. hubrichti populations. The results of this effort will be used to rank pixels from most to least beneficial, then identify land owners in the most useful areas for salamanders who are willing to sell or enter into a permanent easement agreement.

Alabama

Post-independence fledgling ecology in a migratory songbird: Implications for breeding-grounds conservation

For migratory songbirds, breeding-grounds conservation and management plans are generally focused on habitat associated with locations of singing males and sometimes nesting females. However, habitat structure is often different in areas used for raising fledglings compared with areas used for song territories, and very little is known about habitat use by fledglings after independence from adult care. From 2010 to 2012, we used radiotelemetry to monitor 68 fledgling golden-winged warblers V ermivora chrysoptera after independence from adult care in mixed managed forests of Minnesota, US and Manitoba, Canada. This species is of high conservation concern in the US, is listed as threatened in Canada and is listed as near threatened on the International Union for Conservation of Nature Red List. We assessed distance and orientation of independent fledgling movements and we used compositional analysis to test for selection among cover types. Fledglings of this species, commonly described as a shrubland specialist, selected mature forest (78% of locations) over all other cover types, and foraged in forest canopy and understory in mixed-species flocks. Fledgling golden-winged warbler movements were apparently associated with habitat optimization (although prioritizing foraging over predator avoidance), and likely not with commencement of migration, or scouting future breeding territories. Ten days after independence, fledglings were an average of 1238&thinsp;m north of their nest, which may be related to homing-target formation and the species' northward range expansion. We conclude that consideration for independent fledgling habitat associations is necessary for developing full-fledged forest management plans on the breeding grounds of migratory songbirds.

Manitoba, Minnesota

A demographic projection model to support conservation decision making for an endangered snake with limited monitoring data

Conservation planning for rare and threatened species is often made more difficult by a lack of research and monitoring data. In such cases, managers may rely on qualitative assessments of species risk that lack explicit acknowledgement of uncertainty. Snakes are a group of conservation concern that are also notoriously difficult to monitor. Here, we demonstrate a quantitative population projection for a data-deficient species, the Puerto Rican boa ( Chilabothrus inornatus ) using expert knowledge and published information about species life history and threats to persistence. Using this model, we simulated population dynamics over 30 years under four scenarios of future urbanization and found that there was an increased probability of population decline as urbanization rates increased. We conduct a sensitivity analysis to evaluate the sensitivity of outcomes to model inputs, a practice that may also be useful in recovery planning. The sensitivity analyses also provide insight into how the future trajectories would change if the elicited demographic rates are incorrect. Even when data are sparse, quantitative methods can often be used to produce rigorous and reproducible estimates of future status with quantifiable uncertainty.

Puerto Rico

Conservation genetics of the endangered Shenandoah salamander (Plethodon shenandoah, Plethodontidae)

The Shenandoah salamander (Plethodon shenandoah) is restricted to three isolated talus outcrops in Shenandoah National Park, VA, USA and has one of the smallest ranges of any tetrapod vertebrate. This species was listed as endangered under the US Endangered Species Act in 1989 over concern that direct competition with the red-backed salamander (Plethodon cinereus), successional habitat changes, and human impacts may cause its decline and possible extinction. We address two issues herein: (1) whether extensive introgression (through long-term hybridization) is present between the two species and threatens the survival of P. shenandoah, and (2) the level of population structure within P. shenandoah. We provide evidence from mtDNA haplotypes that shows no genetic differentiation among the three isolates of P. shenandoah, suggesting that their fragmentation is a geologically recent event, and/or that the isolates are still connected by occasional gene flow. There is also no evidence for extensive introgression of alleles in either direction between P. cinereus and P. shenandoah, which suggests that P. shenandoah may not be in danger of being genetically swamped out through hybridization with P. cinereus.

Animal Conservation

Forecasting dynamics of a recolonizing wolf population under different management strategies

Species recovery can be influenced by a wide variety of factors, such that predicting the spatiotemporal dynamics of recovering species can be exceedingly difficult. These predictions, however, are valuable for decision makers tasked with managing species and determining their legal status. We applied a spatially explicit projection model to estimate population growth and viability of gray wolves ( Canis lupus ) from 2021 to 2070 in the state of Washington, USA, where wolves have been naturally recolonizing since the establishment of the first resident pack in 2008. Using this model, we predicted the effects of 12 scenarios relating to management actions (e.g., lethal removals by the state agency, translocation, recreational harvest) and system uncertainties (e.g., immigration from out of state, disease) on the probability of meeting Washington's wolf recovery goals, the probability of extinction, and other metrics related to population status. Population recovery was defined under Washington's Wolf Conservation and Management Plan as four breeding pairs in each of three recovery regions and three additional breeding pairs anywhere in the state. The baseline and two translocation scenarios indicated a high (> 90%) probability of wolf recovery in Washington by 2070, but scenarios related to harvest mortality (removal of 5% of the population every 6 months), increased lethal removals (removal of 8.53% of the population across the state each year), and cessation of immigration from out of state resulted in probabilities of < 0.20 (0.01, 0.04, and 0.17, respectively) of meeting recovery goals by 2070. Only two scenarios of 12 (increased harvest and lethal removals scenarios) resulted in a geometric mean of population growth ≤ 1, indicating long-term population stability or growth for most scenarios. Our results suggest that wolves will continue to recolonize Washington and that recovery goals are likely to be met so long as harvest and lethal removals are not at unsustainable levels and adjacent populations support immigration into Washington.

Washington

Low genetic variation and evidence of limited dispersal in the regionally important Belize manatee

The Antillean subspecies of the West Indian manatee Trichechus manatus is found throughout Central and South America and the Caribbean. Because of severe hunting pressure during the 17th through 19th centuries, only small populations of the once widespread aquatic mammal remain. Fortunately, protections in Belize reduced hunting in the 1930s and allowed the country's manatee population to become the largest breeding population in the Wider Caribbean. However, increasing and emerging anthropogenic threats such as coastal development, pollution, watercraft collision and net entanglement represent challenges to this ecologically important population. To inform conservation and management decisions, a comprehensive molecular investigation of the genetic diversity, relatedness and population structure of the Belize manatee population was conducted using mitochondrial and microsatellite DNA. Compared with other mammal populations, a low degree of genetic diversity was detected ( H E =0.455; N A =3.4), corresponding to the small population size and long-term exploitation. Manatees from the Belize City Cayes and Southern Lagoon system were genetically different, with microsatellite and mitochondrial F ST values of 0.029 and 0.078, respectively ( P ≤0.05). This, along with the distinct habitats and threats, indicates that separate protection of these two groups would best preserve the region's diversity. The Belize population and Florida subspecies appear to be unrelated with microsatellite and mitochondrial F ST values of 0.141 and 0.63, respectively ( P ≤0.001), supporting the subspecies designations and suggesting low vagility throughout the northern Caribbean habitat. Further monitoring and protection may allow an increase in the Belize manatee genetic diversity and population size. A large and expanding Belize population could potentially assist in the recovery of other threatened or functionally extinct Central American Antillean manatee populations.

Animal Conservation

Using periodic matrix models to simulate the effectiveness of alternative reintroduction strategies for lizards on a seasonal tropical island

Conservation translocations and reintroductions are widely used to improve conservation outcomes for declining species. Reintroductions are unlikely to be successful if the threats that led to the extirpation of the focal species, such as non-native predators, have not been ameliorated. The non-native brown treesnake ( Boiga irregularis ) was introduced to Guam in the mid-20th century and has devastated the native lizard fauna of this Pacific Island. Native lizard populations persist on small islands near Guam and could act as sources for reintroductions. Recently, trapping methods have been shown to greatly reduce the density of brown treesnakes, raising the possibility that native lizards could be reintroduced where brown treesnake abundance is controlled. Here we developed demographic models for four lizard species present on Guam to assess their validity as a trial for reintroducing species to meet native species recovery goals. These four species vary in their size, activity, and susceptibility to predation by brown treesnakes. We used periodic matrix models to simulate changes in lizard vital rates driven by the wet-dry seasonal cycle found on Guam. We found that the release of 60 individuals of each species was likely to result in successful reintroduction outcomes provided adult survival in the wild is similar to rates estimated based on life history parameters. Demographic models can be used to forecast reintroduction outcomes, including predicting the probability of success and evaluating causes of failure in the event that populations do not establish.

Guam

Estimating distribution and connectivity of recolonizing American marten in the northeastern United States using expert elicitation techniques

The American marten Martes americana is a species of conservation concern in the northeastern United States due to widespread declines from over‐harvesting and habitat loss. Little information exists on current marten distribution and how landscape characteristics shape patterns of occupancy across the region, which could help develop effective recovery strategies. The rarity of marten and lack of historical distribution records are also problematic for region‐wide conservation planning. Expert opinion can provide a source of information for estimating species–landscape relationships and is especially useful when empirical data are sparse. We created a survey to elicit expert opinion and build a model that describes marten occupancy in the northeastern United States as a function of landscape conditions. We elicited opinions from 18 marten experts that included wildlife managers, trappers and researchers. Each expert estimated occupancy probability at 30 sites in their geographic region of expertise. We, then, fit the response data with a set of 58 models that incorporated the effects of covariates related to forest characteristics, climate, anthropogenic impacts and competition at two spatial scales (1.5 and 5 km radii), and used model selection techniques to determine the best model in the set. Three top models had strong empirical support, which we model averaged based on AIC weights. The final model included effects of five covariates at the 5‐km scale: percent canopy cover (positive), percent spruce‐fir land cover (positive), winter temperature (negative), elevation (positive) and road density (negative). A receiver operating characteristic curve indicated that the model performed well based on recent occurrence records. We mapped distribution across the region and used circuit theory to estimate movement corridors between isolated core populations. The results demonstrate the effectiveness of expert‐opinion data at modeling occupancy for rare species and provide tools for planning marten recovery in the northeastern United States.

Animal Conservation

Survey design optimization for monitoring wildlife communities in areas managed for federally endangered species

In wildlife communities composed of federally endangered species, there are often several species of conservation concern that have not yet warranted federally mandated protection. These species often need continued monitoring to inform the direction of future management. While recovering endangered species is an important conservation goal, practitioners are challenged by balancing federally mandated protocols with actions that promote non-listed priority species. Practitioners need an understanding of how focused, single-species management actions may affect non-listed priority species, but developing a monitoring protocol that can detect such effects with limited resources is a challenge. Here we use constrained optimization as a path to identifying a sampling scheme that overcomes these logistical challenges and then illustrate its potential in the Sandhills region of North Carolina, USA. Using empirical results from multi-year avian community monitoring, we parameterized simulations to understand how varying the number of sampling locations and site visits affected the optimal monitoring protocol across three different avian community composition scenarios: a community with (1) 10 percent, (2) 25 percent, or (3) 50 percent non-listed priority species. We found the greatest rate of change in precision of community-level metrics such as species richness by increasing sampling replicates when surveying up to 50 sites. Importantly, this trend was apparent across all three community scenarios, indicating relatively predictable changes in uncertainty regardless of community composition. In contrast, increasing the sampling frequency did not consistently reduce uncertainty in species-level parameters such as occupancy probability. Concerningly, we saw the greatest variation when communities were comprised of 50 percent non-listed species suggesting increasingly complex monitoring protocols may be required if the number of non-listed priority species continues to increase. Practitioners could consider reducing detection error of priority species through increasing sampling frequency, as this can strongly affect optimization study designs.

North Carolina