Evidence for regular sporulation by Haplosporidium nelsoni (MSX) (Ascetospora; Haplosporidiidae) in spat of the American oyster, Crassostrea virginica.

The spore stage of Haplosporidium nelsoni, the ascetosporan parasite causing multinucleated sphere unknown (MSX) disease in oysters, Crassostrea virginica, has been reported so rarely (less than 0.01% of infected oysters) that a second host has been postulated. However, recent intensive sampling of young (less than 1 year) oysters in Delaware Bay, U.S. suggests that spore formation occurs regularly in this group and that spores are produced in at least 75-85% of all infections reaching the advanced stage. Sporulation was seasonal, occurring over two to three weeks in late June/early July and again in late summer/early fall. Our data indicate that sporulation by H. nelsoni in oysters is more common than previously suspected, occurring in a segment of the host population that may not have been sufficiently sampled in the past, and that a direct life cycle should be reconsidered.     

STATUS OF THE PACIFIC WALRUS POPULATION, 1950–19891

The Pacific walrus (Odobenus rosmarus divergens) population is an important ecological and economic resource of the Bering Sea region. We describe population change, beginning with a low in 1950, through a high in about 1980, and ending in 1989. Estimates of abundance for the years after 1989 were not attempted due to the lack of harvest data and other population parameters. Selective hunting practices resulted in biased data regarding population composition and reproductive performance. Rates of reproduction had to be estimated from ovarian data, which indicated a dramatic drop in the 1980s. High harvests in the 1980s likely contributed to a decline in the population, but uncertainties as to accuracy of population estimates and other data raise reasonable doubts, especially with respect to the number of males, for which the most recent (1985) population estimate suggests a sharp decline. Past population estimates were revised upwards to compensate for walruses underwater and not seen in aerial surveys. The weaknesses in the available data make it clear that effective management of the population will require many improvements in collection of data regarding harvests, population structure, reproduction, and population trend.     

Evidence for Regular Sporulation by Haplosporidium nelsoni (MSX) (Ascetospora; Haplosporidiidae) in Spat of the American Oyster, Crassostrea virginica

The spore stage of Haplosporidium nelsoni , the ascetosporan parasite causing multinucleated sphere unknown (MSX) disease in oysters, Crassostrea virginica , has been reported so rarely (≥0.01% of infected oysters) that a second host has been postulated. However, recent intensive sampling of young (≥1 year) oysters in Delaware Bay, U.S. suggests that spore formation occurs regularly in this group and that spores are produced in at least 75–85% of all infections reaching the advanced stage. Sporulation was seasonal, occurring over two to three weeks in late June/early July and again in late summer/early fall. Our data indicate that sporulation by H. nelsoni in oysters is more common than previously suspected, occurring in a segment of the host population that may not have been sufficiently sampled in the past, and that a direct life cycle should be reconsidered.     

AERIAL CENSUS OF PACIFIC WALRUSES IN THE CHUKCHI SEA, 1985

The U.S. Fish and Wildlife Service conducted a survey of the walruses in the pack ice of the Chukchi Sea between 16 September and 2 October 1985, as part of a joint effort with the Soviet Union to estimate the size of the Pacific walrus population. American observers conducted censuses from two aircraft along randomly selected north–south lines over the pack ice. The observers counted walruses within a constant viewing angle that corresponded to a total strip width of 1.38 km at an altitude of 152 m.
In nine days of flying, 15,312 walruses were observed, of which 10,140 were on 5,764 km² of census strips. Few walruses were observed east of 161°W longitude or west of 170°W longitude, hence the census effort was stratified. Walrus concentrations between 161° and 170° shifted slightly westward during the 2-wk duration of the censuses. The differences among days in observed walrus density were due to changes in the numbers of walruses on the ice within 37 km of the ice edge. The number of observable walruses in pack ice of the eastern Chukchi Sea was estimated to be 62,177 (SD = 19,480), based on censuses conducted on 29 and 30 September and 1 October. At that time there were also at least 15,238 in Bristol Bay, Bering Sea. The Soviets counted 39,572 on the shores of the western Chukchi and Bering seas and estimated 115,531 in pack ice of the western Chukchi Sea. Summing U.S. and Soviet estimates, the total population of Pacific walruses in 1985 was 232,518. This number was comparable with earlier estimates from censuses conducted jointly by the U.S. and the Soviets. However, information on fraction hauled out, segregation, and movements is needed for more precise estimates.     

Demography of the Pacific walrus (Odobenus rosmarus divergens) in a changing Arctic

The Pacific walrus (Odobenus rosmarus divergens) is a candidate to be listed as an endangered species under United States law, in part, because of climate change‐related concerns. While the population was known to be declining in the 1980s and 1990s, its recent status has not been determined. We developed Bayesian models of walrus population dynamics to assess the population by synthesizing information on population sizes, age structures, reproductive rates, and harvests for 1974–2015. Candidate models allowed for temporal variation in some or all vital rates, as well as density dependence or density independence in reproduction and calf survival. All selected models indicated that the population underwent a multidecade decline, which began moderating in the 1990s, and that annual reproductive rate and natural calf survival rates rose over time in a density‐dependent manner. However, selected models were equivocal regarding whether the natural juvenile survival rate was constant or decreasing over time. Depending on whether juvenile survival decreased after 1998, the population growth rate either increased during 1999–2015 or stabilized at a lesser level of decline than seen in the 1980s. The probability that the population was still declining in 2015 ranged from 45% to 87%.     

TRENDS IN AGE STRUCTURE AND PRODUCTIVITY OF PACIFIC WALRUSES HARVESTED IN THE BERING STRAIT REGION OF ALASKA, 1952–2002

Pacific walruses ( Odobenus rosmarus divergens ) are harvested by subsistence hunters in Alaska as they migrate north through the Bering Strait in the spring. Harvest records and biological specimens have been collected from the Bering Strait communities of Little Diomede, Gambell, and Savoonga since the 1950s. Harvest levels in the Bering Strait region peaked in the late 1980s and declined thereafter; however, there was considerable variation in the size and composition of the harvests among communities and over time. The relationships among characteristics of the community harvests and the presence of temporal trends were investigated using generalized linear models. The proportion of females in the catch increased over time in all three communities, while the proportion pregnant among harvested females declined over the range of sample years. The ages of harvested walruses increased over time in all three communities through the 1980s, after which trends in age stabilized or began to decline. The age of first reproduction was significantly older for females sampled in 1975–1985 than for females sampled between 1952 and 1962 or 1992 and 1998. Factors thought to have influenced the size and composition of the catch over the past 50 yr include hunter preferences, harvest management regimes, environmental conditions, and changes in the population itself.     

Modelling animal populations in changing landscapes

Models of Mobile Animal Populations (MAP models) simulate long-term land use changes, population trends and patterns of biological diversity on landscapes of 10³-10⁵ ha. MAP models can incorporate information about past land-use patterns and management practices and can project future patterns based on management plans. We illustrate this approach with an example of how implementation of a U.S. Forest Service management plan at the Savannah River Site in South Carolina, U.S.A., might influence population trends of Bachman's Sparrow Aimophila aestivalis, a relatively rare and declining species in southeastern pine forests. In this case, a management plan, largely designed to improve conditions for an endangered species, Red-cockaded Woodpecker Picoides borealis, may have a negative impact, at least in the short term, on another species of management concern, Bachman's Sparrow.
In a parallel processing version of the MAP models, a single landscape that would ordinarily be too large or detailed to be simulated on a single computer is subdivided into a number of smaller landscapes, and each landscape is simulated in parallel, either on a single multi-tasking machine or on a group of networked machines. With this approach we are attempting to determine just how large a landscape must be before the dynamics of a population within it are more or less independent of factors beyond the landscape boundaries.     

Indication of two Pacific walrus stocks from whole tooth elemental analysis

The Pacific walrus (Odobenus rosmarus divergens) is considered to be a single panmictic population for management purposes. However, studies on population structuring in this species are limited; in part, because portions of the population’s range are often inaccessible. Therefore, alternative and complementary methods for investigating stock structure in the Pacific walrus are of particular interest. We used measures of elemental concentrations in whole tooth sections from ICP-MS in a discriminant analysis to investigate evidence of stock separation between walruses from two of three known breeding areas (S.E. Bering, St Lawrence, and Anadyr Gulf). Elemental compositions of teeth from female and male walruses from the S.E. Bering and St Lawrence breeding areas were significantly different, providing evidence of separate stocks. We also obtained insights into the potential relation of walruses from non-breeding areas to walruses from these breeding groups based on similarities in their dental elemental profiles.     

Climate Change and Global Health: Quantifying a Growing Ethical Crisis

Climate change, as an environmental hazard operating at the global scale, poses a unique and “involuntary exposure” to many societies, and therefore represents possibly the largest health inequity of our time. According to statistics from the World Health Organization (WHO), regions or populations already experiencing the most increase in diseases attributable to temperature rise in the past 30 years ironically contain those populations least responsible for causing greenhouse gas warming of the planet. Average global carbon emissions approximate one metric ton per year (tC/yr) per person. In 2004, United States per capita emissions neared 6 tC/yr (with Canada and Australia not far behind), and Japan and Western European countries range from 2 to 5 tC/yr per capita. Yet developing countries’ per capita emissions approximate 0.6 tC/yr, and more than 50 countries are below 0.2 tC/yr (or 30-fold less than an average American). This imbalance between populations suffering from an increase in climate-sensitive diseases versus those nations producing greenhouse gases that cause global warming can be quantified using a “natural debt” index, which is the cumulative depleted CO₂ emissions per capita. This is a better representation of the responsibility for current warming than a single year’s emissions. By this measure, for example, the relative responsibilities of the U.S. in relation to those of India or China is nearly double that using an index of current emissions, although it does not greatly change the relationship between India and China. Rich countries like the U.S. have caused much more of today’s warming than poor ones, which have not been emitting at significant levels for many years yet, no matter what current emissions indicate. Along with taking necessary measures to reduce the extent of global warming and the associated impacts, society also needs to pursue equitable solutions that first protect the most vulnerable population groups; be they defined by demographics, income, or location. For example, according to the WHO, 88% of the disease burden attributable to climate change afflicts children under age 5 (obviously an innocent and “nonconsenting” segment of the population), presenting another major axis of inequity. Not only is the health burden from climate change itself greatest among the world’s poor, but some of the major mitigation approaches to reduce the degree of warming may produce negative side effects disproportionately among the poor, for example, competition for land from biofuels creating pressure on food prices. Of course, in today’s globalized world, eventually all nations will share some risk, but underserved populations will suffer first and most strongly from climate change. Moreover, growing recognition that society faces a nonlinear and potentially irreversible threat has deep ethical implications about humanity’s stewardship of the planet that affect both rich and poor.     

CALCULATING LIMITS TO THE ALLOWABLE HUMAN-CAUSED MORTALITY OF CETACEANS AND PINNIPEDS

A simulation method was developed for identifying populations with levels of human-caused mortality that could lead to depletion, taking into account the uncertainty of available information. A mortality limit (termed the Potential Biological Removal, PBR , under the U. S. Marine Mammal Protection Act) was calculated as the product of a minimum population estimate ( N _MIN), one-half of the maximum net productivity rate ( R _MAX), and a recovery factor ( F _R). Mortality limits were evaluated based on whether at least 95% of the simulated populations met two criteria: (1) that populations starting at the maximum net productivity level ( MNPL ) stayed there or above after 20 yr, and (2) that populations starting at 30% of carrying-capacity ( K ) recovered to at least MNPL after 100 yr. Simulations of populations that experienced mortality equal to the PBR indicated that using approximately the 20th percentile (the lower 60% log-normal confidence limit) of the abundance estimate for N _MIN met the criteria for both cetaceans (assuming R _MAX= 0.04) and pinnipeds (assuming R _MAX= 0.12). Additional simulations that included plausible levels of bias in the available information indicated that using a value of 0.5 for F _R would meet both criteria during these "bias trials." It is concluded that any marine mammal population with an estimate of human-caused mortality that is greater than its PBR has a level of mortality that could lead to the depletion of the population. The simulation methods were also used to show how mortality limits could be calculated to meet conservation goals other than the U. S. goal of maintaining populations above MNPL .     

Pacific Walrus and climate change: observations and predictions

The extent and duration of sea-ice habitats used by Pacific walrus (Odobenus rosmarus divergens) are diminishing resulting in altered walrus behavior, mortality, and distribution. I document changes that have occurred over the past several decades and make predictions to the end of the 21st century. Climate models project that sea ice will monotonically decline resulting in more ice-free summers of longer duration. Several stressors that may impact walruses are directly influenced by sea ice. How these stressors materialize were modeled as most likely-case, worst-case, and best-case scenarios for the mid- and late-21st century, resulting in four comprehensive working hypotheses that can help identify and prioritize management and research projects, identify comprehensive mitigation actions, and guide monitoring programs to track future developments and adjust programs as needed. In the short term, the most plausible hypotheses predict a continuing northward shift in walrus distribution, increasing use of coastal haulouts in summer and fall, and a population reduction set by the carrying capacity of the near shore environment and subsistence hunting. Alternatively, under worst-case conditions, the population will decline to a level where the probability of extinction is high. In the long term, walrus may seasonally abandon the Bering and Chukchi Seas for sea-ice refugia to the northwest and northeast, ocean warming and pH decline alter walrus food resources, and subsistence hunting exacerbates a large population decline. However, conditions that reverse current trends in sea ice loss cannot be ruled out. Which hypothesis comes to fruition depends on how the stressors develop and the success of mitigation measures. Best-case scenarios indicate that successful mitigation of unsustainable harvests and terrestrial haulout-related mortalities can be effective. Management and research should focus on monitoring, elucidating effects, and mitigation, while ultimately, reductions in greenhouse gas emissions are needed to reduce sea-ice habitat losses.     

OBSERVER VARIABILITY IN PINNIPED COUNTS: GROUND-BASED ENUMERATION OF WALRUSES AT HAUL-OUT SITES

Pinnipeds are often monitored by counting individuals at haul-out sites, but the often large numbers of densely packed individuals at these sites are difficult to enumerate accurately. Errors in enumeration can induce bias and reduce precision in estimates of population size and trend. We used data from paired observers monitoring walrus haul-outs in Bristol Bay, Alaska, to quantify observer variability and assess its relative importance. The probability of a pair of observers making identical counts was < 0.1 for walrus groups with >50 individuals. Mean count differences ranged up to 25% for the largest counts, depending on beach and observers. In at least some cases, there was a clear tendency for counts of one observer to be consistently greater than counts of the other observer in a pair, indicating that counts of at least one of the observers were biased. These results suggest that efforts to improve accuracy of counts will be worthwhile. However, we also found that variation among observers was relatively small compared to variation among visits to a beach so that efforts to account for other sources of variation will be more important.     

Red snapper management in the Gulf of Mexico: science- or faith-based?

The most controversial fishery in U.S. waters of the Gulf of Mexico (Gulf) is for northern red snapper Lutjanus campechanus, which collapsed in the late 1980s when stock biomass became too low to be fished commercially in the eastern Gulf. Red snapper management began in 1989; the stock is now showing signs of recovery. The Gulf of Mexico Fishery Management Council has been slow to sufficiently reduce catches of the directed fisheries to rebuild the stock in a timely fashion, although compliance with the Magnuson-Stevens Reauthorization Act of 2006 (MSRA) required substantial cuts in the harvest of red snapper beginning in 2007. In our opinion, this could have been avoided if conservative management practices had been adopted earlier. We believe that ‘faith-based fisheries’ arguments have been used to defer effective management of red snapper in the Gulf, which in turn has strained the relationship between science, management, and stakeholders there. We provide a simple empirical argument and alternate interpretations of a recently published perspective on the historical fishery of red snapper in the Gulf to conclude that the preponderance of evidence used in the agency stock assessment process, and the simple arguments made here, do not support the perspective that the red snapper stock has increased in size sufficiently to defer compliance with the MSRA.     

Phylogenetic analysis reveals a correlation between the expansion of very virulent infectious bursal disease virus and reassortment of its genome segment B

Infectious bursal disease virus (IBDV) is a birnavirus causing immunosuppressive disease in chickens. Emergence of the very virulent form of IBDV (vvIBDV) in the late 1980s dramatically changed the epidemiology of the disease. In this study, we investigated the phylogenetic origins of its genome segments and estimated the time of emergence of their most recent common ancestors. Moreover, with recently developed coalescence techniques, we reconstructed the past population dynamics of vvIBDV and timed the onset of its expansion to the late 1980s. Our analysis suggests that genome segment A of vvIBDV emerged at least 20 years before its expansion, which argues against the hypothesis that mutation of genome segment A is the major contributing factor in the emergence and expansion of vvIBDV. Alternatively, the phylogeny of genome segment B suggests a possible reassortment event estimated to have taken place around the mid-1980s, which seems to coincide with its expansion within approximately 5 years. We therefore hypothesize that the reassortment of genome segment B initiated vvIBDV expansion in the late 1980s, possibly by enhancing the virulence of the virus synergistically with its existing genome segment A. This report reveals the possible mechanisms leading to the emergence and expansion of vvIBDV, which would certainly provide insights into the scope of surveillance and prevention efforts regarding the disease.     

The United States and the new law of the sea

Abstract

The official U.S. statement on the 32nd meeting of the International Whaling Commission (IWC termed the outcome a “modest success.”; However, from the point of view of conservationists—i.e., governments and private groups seeking greater protection for the world's endangered whales—the results of the 1980 IWC session can be described at best as satisfactory. The disappointing U.S. performance at this and other recent Commission proceedings strongly suggests the need for a reassessment of American cetacean policy and the way in which we pursue this policy in the IWC.

The discussion will begin with a review of U.S. objectives in the IWC in relation to national and international law governing cetaceans. The focus will then turn to the role played by the United States in the Commission's deliberations on selected issues in 1980. The conclusion will set forth recommendations for improvements in American policy and strategy in the IWC.     

Identifying highly connected counties compensates for resource limitations when evaluating national spread of an invasive pathogen

Surveying invasive species can be highly resource intensive, yet near-real-time evaluations of invasion progress are important resources for management planning. In the case of the soybean rust invasion of the United States, a linked monitoring, prediction, and communication network saved U.S. soybean growers approximately $200 M/yr. Modeling of future movement of the pathogen (Phakopsora pachyrhizi) was based on data about current disease locations from an extensive network of sentinel plots. We developed a dynamic network model for U.S. soybean rust epidemics, with counties as nodes and link weights a function of host hectarage and wind speed and direction. We used the network model to compare four strategies for selecting an optimal subset of sentinel plots, listed here in order of increasing performance: random selection, zonal selection (based on more heavily weighting regions nearer the south, where the pathogen overwinters), frequency-based selection (based on how frequently the county had been infected in the past), and frequency-based selection weighted by the node strength of the sentinel plot in the network model. When dynamic network properties such as node strength are characterized for invasive species, this information can be used to reduce the resources necessary to survey and predict invasion progress.     

WALRUS LIFE-HISTORY MOVEMENTS RECONSTRUCTED FROM LEAD ISOTOPES IN ANNUAL LAYERS OF TEETH

Putative life history movements of male Atlantic walrus ( Odobenus rosmarus nsmarus ) harvested at Sanirajak in northern Foxe Basin, Canada, were reconstructed by comparing Pb isotopes in tooth cementum growth layer groups (GLGs) to those in whole-cementum samples from other walrus of Foxe Basin and nearby areas. The analyses yielded three life history patterns including one, "prodigal sons," not previously recognized in large mammals. Isotope ratios in GLGs of "local" Sanirajak walrus were relatively constant over time and similar to the whole-cementum data for most walrus caught in northern Foxe Basin. An "immigrant" walrus acquired the local signal only in young adulthood. In two "prodigal sons," GLG Pb isotope ratios changed significantly from the local signature after weaning but returned to it in late maturity, shortly before being harvested. Possible recruitment and dispersal areas for the two types of outliers included regions in northern and eastern Hudson Bay, some 300–600 km distant. The return of mature males to their natal group after associating with other groups for several years, as seen in some of the walrus, may be more common among mammals than previously thought. However, without a reconsttuctive technique such as employed here, it could only be detected through long-term studies of known individuals and all the relevant social groups.     

The Laptev Sea walrus Odobenus rosmarus laptevi: an enigma revisited

The walrus ( Odobenus rosmarus ) is in some current systematic schemes divided into three subspecies: O. r. rosmarus in the North Atlantic, O.   r. divergens in the North Pacific and O.   r. laptevi in the Laptev Sea. These three subspecies have been described as differing in body size, but the taxonomic status of O.   r. laptevi is disputed. The current study applies molecular and morphometric methods to assess the taxonomic status of O.   r. laptevi and to analyse the systematic and phylogeographic relationships between the three purported walrus subspecies. Tusk length and tusk circumference were measured from the few skulls available of O.   r. laptevi , and the obtained values were within the ranges reported for Pacific walruses. Thus, morphologically, subspecies status for O.   r. laptevi is not supported according to the Amadon–Mayr '75% rule'. Phylogenetic analyses and haplotype networks based on mitochondrial nucleotide sequence data of NADH dehydrogenase 1, 16S rRNA, cytochrome oxidase I and the d -loop of the control region of the historic O.   r. laptevi bone material and contemporary O.   r. rosmarus and O.   r. divergens showed that the Laptev Sea walrus groups with individuals from the North Pacific. Thus, the mitochondrial sequence data do not support the recognition of three walrus subspecies as reciprocally monophyletic evolutionary units with independent evolutionary histories. Only O.   r. rosmarus and O.   r. divergens meet this criterion with the present sampling. Accordingly, we recommend that Odobenus r. laptevi be abandoned and the Laptev walrus instead be recognized as the westernmost population of the Pacific walrus, Odobenus r. divergens. However, further research is recommended to assess whether the Laptev walrus could be considered as a significant unit in terms of conservation and management, since it is unique in several ecological parameters.     

The founding of a southern elephant seal colony

The only large mainland colony of southern elephant seals (Mirounga leonina) is on Península Valdés, at 42°S, in Argentine Patagonia. Censuses of pups have been carried out regularly there since 1970, and the population grew five‐fold by 2010. Here we use Bayesian modeling tools to make rigorous estimates of the rate of population growth, r, and to estimate survival and recruitment parameters that could account for the growth, incorporating observation error across different census methods. In the 1970s, r= 8%/yr, but has slowed to <1%/yr over the past decade. Using explicit demographic models, we established that the high growth of the 1970s was consistent with adult and juvenile survival at the upper end of published values (0.87/yr adult female survival; 0.40 juvenile survivorship to age four); the decline in the rate of population growth from 1970 to 2010 can be described by density‐dependent reductions in adult and juvenile survival that fall well within published variation. Extrapolating empirical models of population growth rate backwards illustrates that the population could have been an established colony, with 100 pups born per year, between 1915 and 1945, consistent with qualitative observations prior to 1950. We conclude that the Valdés colony was founded by a few immigrants early in the 20th century and has been growing mostly by internal recruitment, with unknown density‐dependent processes causing a reduction in growth and stabilization at 15,000–16,000 pups born.     

RISK OF EXTIRPATION OF STELLER SEA LIONS IN THE GULF OF ALASKA AND ALEUTIAN ISLANDS: A POPULATION VIABILITY ANALYSIS BASED ON ALTERNATIVE HYPOTHESES FOR WHY SEA LIONS DECLINED IN WESTERN ALASKA

We estimated the risk that the Steller sea lion will be extirpated in western Alaska using a population viability analysis (PVA) that combined simulations with statistically fitted models of historical population dynamics. Our analysis considered the roles that density‐dependent and density‐independent factors may have played in the past, and how they might influence future population dynamics. It also established functional relationships between population size, population growth rate and the risk of extinction under alternative hypotheses about population regulation and environmental variability. These functional relationships can be used to develop recovery criteria and guide research and management decisions. Life table parameters (e.g., birth and survival rates) operating during the population decline (1978–2002) were estimated by fitting simple age‐structured models to time‐series of pup and non‐pup counts from 33 rookeries (subpopulations). The PVA was carried out by projecting all 33 subpopulations into the future using these estimated site‐specific life tables (with associated uncertainties) and different assumptions about carrying capacities and the presence or absence of density‐dependent population regulation. Results suggest that the overall predicted risk of extirpation of Steller sea lions as a species in western Alaska was low in the next 100 yr under all scenarios explored. However, most subpopulations of Steller sea lions had high probabilities of going extinct within the next 100 yr if trends observed during the 1990s were to continue. Two clusters of contiguous subpopulations occurring in the Unimak Pass area in the western Gulf of Alaska/eastern Aleutian Islands and the Seguam–Adak region in the central Aleutian Islands had relatively lower risks of extinction. Risks of extinction for a number of subpopulations in the Gulf of Alaska were reduced if the increases observed since the late 1990s continue into the future. The risks of subpopulations going extinct were small when density‐dependent compensation in birth and survival rates was assumed, even when random stochasticity in these vital rates was introduced.