Daniel H. Monson

UNDERSTANDING AND PREDICTING ECOLOGICAL DYNAMICS: ARE MAJOR SURPRISES INEVITABLE

Ecological surprises, substantial and unanticipated changes in the abundance of one or more species that result from previously unsuspected processes, are a common outcome of both experiments and observations in community and population ecology. Here, we give examples of such surprises along with the results of a survey of well-established field ecologists, most of whom have encountered one or more surprises over the course of their careers. Truly surprising results are common enough to require their consideration in any reasonable effort to characterize nature and manage natural resources. We classify surprises as dynamic-, pattern-, or intervention-based, and we speculate on the common processes that cause ecological systems to so often surprise us. A long-standing and still growing concern in the ecological literature is how best to make predictions of future population and community dynamics. Although most work on this subject involves statistical aspects of data analysis and modeling, the frequency and nature of ecological surprises imply that uncertainty cannot be easily tamed through improved analytical procedures, and that prudent management of both exploited and conserved communities will require precautionary and adaptive management approaches.

Could residual oil from the Exxon Valdez spill create a long-term population ''sink'' for sea otters in Alaska?

Over 20 years ago, the Exxon Valdez oil tanker spilled 42 million L of crude oil into the waters of Prince William Sound, Alaska, USA. At the time of the spill, the sea otter (Enhydra lutris) population inhabiting the spill area suffered substantial acute injuries and loss. Subsequent research has resulted in one of the best-studied species responses to an oil spill in history. However, the question remains: Is the spill still influencing the Prince William Sound sea otter population? Here we fit time-varying population models to data for the sea otter population of western Prince William Sound to quantify the duration and extent of mortality effects from the spill. We hypothesize that the patchy nature of residual oil left in the environment has created a source-sink population dynamic. We fit models using the age distributions of both living and dying animals and estimates of sea otter population size to predict the number of sea otters in the hypothesized sink population and the number lost to this sink due to chronic exposure to residual oil. Our results suggest that the sink population has remained at just over 900 individuals (95% CI: 606-960) between 1990 and 2009, during which time prime-age survival remained 2-6% below pre-spill levels. This reduced survival led to chronic losses of ;900 animals over the past two decades, which is similar in magnitude to the number of sea otter deaths documented in western Prince William Sound during the acute phase of the spill. However, the unaffected source population appears to be counterbalancing these losses, with the model indicating that the sea otter population increased from ;2150 individuals in 1990 to nearly 3000 in 2009. The most optimistic interpretation of our results suggests that mortality effects dissipated between 2005 and 2007. Our results suggest that residual oil can affect wildlife populations on time scales much longer than previously believed and that cumulative chronic effects can be as significant as acute effects. Further, source-sink population dynamics can explain the slow recovery observed in the spill-affected western Prince William Sound sea otter population and are consistent with available data.

CHEMICAL ANESTHESIA OF NORTHERN SEA OTTERS (ENHYDRA LUTRIS): RESULTS OF PAST FIELD STUDIES

Abstract Between 1987 and 1997, we chemically immobilized 597 wild sea otters (Enhydra lutris) in Alaska for the collection of biological samples or for surgical instrumentation. One drug-related sea otter fatality occurred during this time. Fentanyl in combination with diazepam produced consistent, smooth inductions with minimal need for supplemental anesthetics during procedures lasting 30–40 min. Antagonism with naltrexone or naloxone was rapid and complete, although we observed narcotic recycling in sea otters treated with naloxone. For surgical procedures, we recommend a fentanyl target dose of 0.33 mg/kg of body mass and diazepam at 0.11 mg/kg. For nonsurgical biological sample collection procedures, we recommend fentanyl at 0.22 mg/kg and diazepam at 0.07 mg/kg. We advise the use of the opioid antagonist naltrexone at a ratio of 2:1 to the total fentanyl administered during processing.

Activity Budgets Derived From Time–Depth Recorders in a Diving Mammal

Abstract We describe a method to convert continuously collected time–depth data from archival time–depth recorders (TDRs) into activity budgets for a benthic-foraging marine mammal. We used data from 14 TDRs to estimate activity-specific time budgets in sea otters (Enhydra lutris) residing near Cross Sound, southeast Alaska, USA. From the TDRs we constructed a continuous record of behavior for each individual over 39–46 days during summer of 1999. Behaviors were classified as foraging (diving to the bottom), other diving (traveling, grooming, interacting), and nondiving (assumed resting). The overall average activity budget (proportion of 24-hr/d) was 0.37 foraging (8.9 hr/d), 0.11 in other diving (2.6 hr/d), and 0.52 nondiving time (12.5 hr/d). We detected significant differences in activity budgets among individuals and between groups within our sample. Historically, the sea otter population in our study area had been expanding and sequentially reoccupying vacant habitat since their reintroduction to the area in the 1960s, and our study animals resided in 2 adjacent yet distinct locations. Males (n = 5) and individuals residing in recently occupied habitat (n = 4) spent 0.28–0.30 of their time foraging (6.7–7.2 hr/d), 0.17–0.18 of their time in other diving behaviors (4.1–4.3 hr/d), and 0.53–0.54 of their time resting (12.7–13.0 hr/d). In contrast, females (n = 9) and individuals residing in longer occupied habitat (n = 10) spent 0.40 of their time foraging (9.6 hr/d), 0.08–0.09 of their time in other diving behaviors (1.9–2.2 hr/d), and 0.51–0.52 of their time resting (12.2–12.5 hr/d). Consistent with these differences, sea otters residing in more recently occupied habitat captured more and larger clams (Saxidomus spp., Protothaca spp., Macoma spp., Mya spp., Clinocardium spp.) and other prey, and intertidal clams were more abundant and larger in this area. We found that TDRs provided data useful for measuring activity time budgets and behavior patterns in a diving mammal over long and continuous time periods. Fortuitous contrasts in time budgets between areas where our study animals resided suggest that activity time budgets estimated from TDRs may be a sensitive indicator of population status, particularly in relation to prey availability.

Assessment of clinical pathology and pathogen exposure in sea otters (Enhydra lutris) bordering the threatened population in Alaska.

Northern sea otter (Enhydra lutris kenyoni) abundance has decreased dramatically over portions of southwest Alaska, USA, since the mid-1980s, and this stock is currently listed as threatened under the Endangered Species Act. In contrast, adjacent populations in south central Alaska, USA, and Russia have been stable to increasing during the same period. Sea otters bordering the area classified in the recent decline were live-captured during 2004–2006 at Bering Island, Russia, and the Kodiak Archipelago, Alaska, USA, to evaluate differences in general health and current exposure status to marine and terrestrial pathogens. Although body condition was lower in animals captured at Bering Island, Russia, than it was at Kodiak, USA, clinical pathology values did not reveal differences in general health between the two regions. Low prevalences of antibodies (>5%) were found in Kodiak, USA, and on Bering Island, Russia, to Toxoplasma gondii, Sarcocystis neurona, and Leptospira interrogans. Exposure to phocine herpesvirus-1 was found in both Kodiak, USA (15.2%), and Bering Island, Russia (2.3%). Antibodies to Brucella spp. were found in 28% of the otters tested on Bering Island, Russia, compared with only 2.7% of the samples from Kodiak, USA. Prevalence of exposure to Phocine distemper virus (PDV) was 41% in Kodiak, USA, but 0% on Bering Island, Russia. Archived sera from southwest and south-central Alaska dating back to 1989 were negative for PDV, indicating exposure occurred in sea otters in Kodiak, USA, in recent years. Because PDV can be highly pathogenic in naïve and susceptible marine mammal populations, tissues should be examined to explore the contribution of this virus to otter deaths. Our results reveal an increase in exposure to pathogens in sea otters in Kodiak, Alaska, USA, since the 1990s.

Estimating Age Ratios and Size of Pacific Walrus Herds on Coastal Haulouts using Video Imaging

During Arctic summers, sea ice provides resting habitat for Pacific walruses as it drifts over foraging areas in the eastern Chukchi Sea. Climate-driven reductions in sea ice have recently created ice-free conditions in the Chukchi Sea by late summer causing walruses to rest at coastal haulouts along the Chukotka and Alaska coasts, which provides an opportunity to study walruses at relatively accessible locations. Walrus age can be determined from the ratio of tusk length to snout dimensions. We evaluated use of images obtained from a gyro-stabilized video system mounted on a helicopter flying at high altitudes (to avoid disturbance) to classify the sex and age of walruses hauled out on Alaska beaches in 2010–2011. We were able to classify 95% of randomly selected individuals to either an 8- or 3-category age class, and we found measurement-based age classifications were more repeatable than visual classifications when using images presenting the correct head profile. Herd density at coastal haulouts averaged 0.88 walruses/m2 (std. err. = 0.02), herd size ranged from 8,300 to 19,400 (CV 0.03–0.06) and we documented ∼30,000 animals along ∼1 km of beach in 2011. Within the herds, dependent walruses (0–2 yr-olds) tended to be located closer to water, and this tendency became more pronounced as the herd spent more time on the beach. Therefore, unbiased estimation of herd age-ratios will require a sampling design that allows for spatial and temporal structuring. In addition, randomly sampling walruses available at the edge of the herd for other purposes (e.g., tagging, biopsying) will not sample walruses with an age structure representative of the herd. Sea ice losses are projected to continue, and population age structure data collected with aerial videography at coastal haulouts may provide demographic information vital to ongoing efforts to understand effects of climate change on this species.

Evaluating the status of individuals and populations: Advantages of multiple approaches and time scales: Chapter 6

The assessment of population status is a central goal of applied wildlife research and essential to the field of wildlife conservation. “Population status” has a number of definitions, the most widely used having to do with the current trajectory of the population (i.e., growing, stable, or declining), or the probability of persistence (i.e., extinction risk), perhaps without any specific knowledge as to the factors driving a population’s dynamics. In contrast, a population’s status relative to the carrying capacity of the environment ( K ) is an ecologically-based definition that explicitly provides information about a major mechanism of population control. That is, it relates to the relative per capita availability of resources to individuals in a population, which can also be used to infer the state of the environment itself. Sea otters in the North Pacific provide an excellent system with which to examine various approaches to assessing population status relative to K . This is because sea otters were nearly extirpated by historic commercial overexploitation in the eighteenth and nineteenth centuries, followed by natural and translocation-aided population recovery during the twentieth century, and this decline and recovery has been relatively well documented. This provided a unique opportunity to study populations at the extremes of the population status spectrum. Here we describe and review the approaches that have been utilized in sea otter research to understand the status of populations relative to resource abundance. Specifically, we will illustrate the utility of various indices of population status for understanding population dynamics using the case study of a second precipitous sea otter decline in the Western Aleutians. The indices or “tools” described here fit into several broad categories including (1) energetic, (2) morphological, and (3) demographic as well as a fourth category of emerging tools that have not yet been employed in many other situations including dietary diversity, community structure, spatial distribution, and gene expression. Overall, a variety of indices used to measure population status throughout the sea otter’s range have provided insights for understanding the mechanisms driving the trajectory of various sea otter populations, which a single index could not, and we suggest using multiple methods to measure a population’s status at multiple spatial and temporal scales. The work described here also illustrates the usefulness of long-term data sets and/or approaches that can be used to assess population status retrospectively, providing information otherwise not available. While not all systems will be as amenable to using all the approaches presented here, we expect innovative researchers could adapt analogous multi-scale methods to a broad range of habitats and species including apex predators occupying the top trophic levels, which are often of conservation concern.

Lactation and resource limitation affect stress responses, thyroid hormones, immune function, and antioxidant capacity of sea otters (Enhydra lutris)

Abstract Lactation is the most energetically demanding stage of reproduction in female mammals. Increased energetic allocation toward current reproduction may result in fitness costs, although the mechanisms underlying these trade‐offs are not well understood. Trade‐offs during lactation may include reduced energetic allocation to cellular maintenance, immune response, and survival and may be influenced by resource limitation. As the smallest marine mammal, sea otters (Enhydra lutris) have the highest mass‐specific metabolic rate necessitating substantial energetic requirements for survival. To provide the increased energy needed for lactation, female sea otters significantly increase foraging effort, especially during late‐lactation. Caloric insufficiency during lactation is reflected in the high numbers of maternal deaths due to End‐Lactation Syndrome in the California subpopulation. We investigated the effects of lactation and resource limitation on maternal stress responses, metabolic regulation, immune function, and antioxidant capacity in two subspecies of wild sea otters (northern: E. l. nereis and southern: E. l. kenyoni) within the California, Washington, and Alaska subpopulations. Lactation and resource limitation were associated with reduced glucocorticoid responses to acute capture stress. Corticosterone release was lower in lactating otters. Cortisol release was lower under resource limitation and suppression during lactation was only evident under resource limitation. Lactation and resource limitation were associated with alterations in thyroid hormones. Immune responses and total antioxidant capacity were not reduced by lactation or resource limitation. Southern sea otters exhibited higher concentrations of antioxidants, immunoglobulins, and thyroid hormones than northern sea otters. These data provide evidence for allocation trade‐offs during reproduction and in response to nutrient limitation but suggest self‐maintenance of immune function and antioxidant defenses despite energetic constraints. Income‐breeding strategists may be especially vulnerable to the consequences of stress and modulation of thyroid function when food resources are insufficient to support successful reproduction and may come at a cost to survival, and thereby influence population trends.