Cystic Echinococcosis in the Arctic and Sub-Arctic

The northern biotype of Echinococcus granulosus occurs throughout the holarctic zones of tundra and taiga, from eastern Fennoscandia to the Bering Strait in Eurasia and in North America from arctic Alaska approximately to the northern border of the United States. The cycle of the cestode is complex in taiga at lower latitudes, because of the greater diversity of potential hosts. In the Arctic and Subarctic, however, four patterns of predator/prey relationships may be discerned. Two natural cycles involve the wolf and wild reindeer and the wolf and elk (moose), respectively. Where deer of the two species coexist, both are prey of the wolf; the interactions of the wolf and elk are here described on the basis of long-term observations made on Isle Royale (in Lake Superior near the southern limit of taiga), where only the wolf and elk serve as hosts for E. granulosus. A synanthropic cycle involving herding-dogs and domesticated reindeer caused hyperendemicity of cystic echinococcosis in arctic Eurasia, mainly in northeastern Siberia. The 4th pattern, a semi-synanthropic cycle, formerly existed in Alaska, wherein sled-dogs of the indigenous hunters became infected by consuming the lungs of wild reindeer. The sequence of changes in life-style inherent in the process of acculturation affected the occurrence of cystic echinococcosis among nomadic Iñupiat in arctic Alaska. When those people became sedentary, the environs of their early villages soon became severely contaminated by feces of dogs, and cases of cystic echinococcosis occurred. Compared to cystic echinococcosis caused by E. granulosus adapted to synanthropic hosts (dog and domestic ungulates), the infection produced by the northern biotype is relatively benign. 0fearly all diagnosed cases of cystic echinococcosis (> 300 in Alaska have occurred in indigenous people; only one fatality has been recorded (in a non-indigenous person). After sled-dogs were replaced by machines, cases have become rare in Alaska. A similar effect has been observed in Fennoscandia, in the Saami and domesticated reindeer. Recent records indicate tbat the prcvalence of cystic echinococcosis is increasing in Russia, suggesting that dogs are used there in herding.

SIDE EFFECTS OF PERSISTENT TOXICANTS

As you can see from the general tenor of the printed program for this seminar, I am in the unenviable position of trying to discourage you from certain types of chemical control; but my assigned topic "Side Effects of Persistent Toxicants," implies that mission. However, my remarks may be somewhat anticlimax at this time, because it is now generally conceded that we need to reevaluate certain chemicals in control work and to restrict or severely curtail use of those that per¬sist for long periods in the environment. So let me detail my reasons for a somewhat negative attitude toward the use of the persistent hydrocarbons from my experience with the effects of these materials on birds. But first a few words of caution about control work in general, which so often disrupts natural processes and leads to new and unforseen difficulties. As an example, I think of the irruption of mice in the Klamath valley in northern California and southern Oregon in the late '50's. Intensive predator control, particularly of coyotes, but also of hawks and owls, was followed by a severe outbreak of mice in the spring of 1958. To combat the plague of mice, poisoned bait (1080 and zinc phosphide) was widely distributed in an area used by 500,000 waterfowl each spring. More than 3,000 geese were poisoned, so driv¬ing parties were organized to keep the geese off the treated fields. Here it seems conceivable that the whole chain of costly events--cost of the original and probably unnecessary predator control, economic loss to crops from the mouse outbreak, another poisoning campaign to combat the mice, loss of valuable waterfowl resources, and man-hours involved in flushing geese from the fields--might have been averted by a policy of not interfering with the original predator-prey relationship. This points to a dilemma we always face. (We create deplorable situations by clumsy interference with natural processes, then seek artificial cures to correct our mistakes.) For example, we spend millions of dollars in seeking cures for cancer, but do little or nothing about restricting the use of known or suspected carcinogens such as nicotine and DDT.

Provisioning Strategies of Antarctic Fur Seals and Chinstrap Penguins Produce Different Responses to Distribution of Common Prey and Habitat

Central-place foragers that must return to a breeding site to deliver food to offspring are faced with trade-offs between prey patch quality and distance from the colony. Among colonial animals, pinnipeds and seabirds may have different provisioning strategies, due to differences in their ability to travel and store energy. We compared the foraging areas of lactating Antarctic fur seals and chinstrap penguins breeding at Seal Island, Antarctica, to investigate whether they responded differently to the distribution of their prey (Antarctic krill and myctophid fish) and spatial heterogeneity in their habitat. Dense krill concentrations occurred in the shelf region near the colony. However, only brooding penguins, which are expected to be time-minimizers because they must return frequently with whole food for their chicks, foraged mainly in this proximal shelf region. Lactating fur seals and incubating penguins, which can make longer trips to increase energy gain per trip, and so are expected to be energy-maximizers, foraged in the more distant (>20 km from the island) slope and oceanic regions. The shelf region was characterized by more abundant, but lower-energy-content immature krill, whereas the slope and oceanic regions had less abundant but higher-energy-content gravid krill, as well as high-energy-content myctophids. Furthermore, krill in the shelf region undertook diurnal vertical migration, whereas those in the slope and oceanic regions stayed near the surface throughout the day, which may enhance the capture rate for visual predators. Therefore, we sug- gest that the energy-maximizers foraged in distant, but potentially more profitable feeding regions, while the time-minimizers foraged in closer, but potentially less profitable regions. Thus, time and energy constraints derived from different provisioning strategies may result in sympatric colonial predator species using different foraging areas, and as a result, some central-place foragers use sub- optimal foraging habitats, in terms of the quality or quantity of available prey.

Monitoring the Prey-Field of Marine Predators: Combining Digital Imaging With Datalogging Tags

There is increasing interest in the diving behavior of marine mammals. However, identifying foraging among recorded dives often requires several assumptions. The simultaneous acquisition of images of the prey encountered, together with records of diving behavior will allow researchers to more fully investigate the nature of subsurface behavior. We tested a novel digital camera linked to a time-depth recorder on Antarctic fur seals (Arctocephalus gazella). During the austral summer 2000-2001, this system was deployed on six lactating female fur seals at Bird Island, South Georgia, each for a single foraging trip. The camera was triggered at depths greater than 10 m. Five deployments recorded still images (640 x 480 pixels) at 3-sec intervals (total 8,288 images), the other recorded movie images at 0.2-sec intervals (total 7,598 frames). Memory limitation (64 MB) restricted sampling to approximately 1.5 d of 5-7 d foraging trips. An average of 8.5% of still pictures (2.4%-11.6%) showed krill (Euphausia superba) distinctly, while at least half the images in each deployment were empty, the remainder containing blurred or indistinct prey. In one deployment krill images were recorded within 2.5 h (16 km, assuming 1.8 m/sec travel speed) of leaving the beach. Five of the six deployments also showed other fur seals foraging in conjunction with the study animal. This system is likely to generate exciting new avenues for interpretation of diving behavior.

Hunting and social behaviour of leopard seals (Hydrurga leptonyx) at Seal Island, South Shetland Islands, Antarctica

The hunting behavior of leopard seals Hydrurga leptonyx was monitored opportunistically at Seal Island, South Shetland Islands, during the austral summers from 1986/87 to 1994/95. Leopard seals used several methods to catch Antarctic fur seal pups Arctocephalus gazella and chinstrap penguins Pygoscelis antarctica, and individuals showed different hunting styles and hunting success. One to two leopard seals per year were responsible for an average of 60% of observed captures of fur seal pups. Leopard seals preyed on penguins throughout the summer, but preyed on fur seal pups only between late December and mid-February. Hunting behavior differed significantly between different locations on the island; fur seals were hunted only at one colony, and penguins were hunted in several areas. The relative abundance of prey types, size of prey in relation to predator, and specialization of individual leopard seals to hunt fur seal prey probably influence individual prey preferences among leopard seals. On five occasions, two leopard seals were seen together on Seal Island. Possible interpretations of the relationship between the interacting leopard seals included a mother-offspring relationship, a consorting male-female pair, and an adult leopard seal followed by an unrelated juvenile. In two incidents at Seal Island, two leopard seals were observed interacting while hunting: one seal captured fur seal pups and appeared to release them to the other seal. Observations of leopard seals interacting during hunting sessions were difficult to confirm as co-operative hunting, but they strongly implied that the two seals were not agonistic toward one another. The hunting success of individual leopard seals pursuing penguins or fur seals is probably high enough for co-operative hunting not to become a common hunting strategy; however, it may occur infrequently when it increases the hunting productivity of the seals.

Population Growth of Antarctic Fur Seals: Limitation by a Top Predator, The Leopard Seal?

Antarctic fur seals (Arctocephalus gazella) in the South Shetland Islands are recovering from 19th-century exploitation more slowly than the main population at South Georgia. To document demographic changes associated with the recovery in the South Shetlands, we monitored fur seal abundance and reproduction in the vicinity of Elephant Island during austral summers from 1986/1987 through 1994/1995. Total births, mean and variance of birth dates, and average daily mortality rates were estimated from daily live pup counts at North Cove (NC) and North Annex (NA) colonies on Seal Island. Sightings of leopard seals (Hydrurga leptonyx) and incidents of leopard seal predation on fur seal pups were recorded opportunistically during daily fur seal research at both sites. High mortality of fur seal pups, attributed to predation by leopard seals frequently observed at NC, caused pup numbers to decline rapidly between January and March (i.e., prior to weaning) each year and probably caused a long-term decline in the size of that colony. The NA colony, where leopard seals were never observed, increased in size during the study. Pup mortality from causes other than leopard seal predation appeared to be similar at the two sites. The number of pups counted at four locations in the Elephant Island vicinity increased slowly, at an annual rate of 3.8%, compared to rates as high as 11% at other locations in the South Shetland Islands. Several lines of circumstantial evidence are consistent with the hypothesis that leopard seal predators limit the growth of the fur seal population in the Elephant Island area and perhaps in the broader population in the South Shetland Islands. The sustained growth of this fur seal population over many decades rules out certain predator–prey models, allowing inference about the interaction between leopard seals and fur seals even though it is less thoroughly studied than predator–prey systems of terrestrial vertebrates of the northern hemisphere. Top-down forces should be included in hypotheses for future research on the factors shaping the recovery of the fur seal population in the South Shetland Islands.

Killer Whales and Marine Mammal Trends in The North Pacific—A Re-Examination of Evidence for Sequential Megafauna Collapse and the Prey-Switching Hypothesis

Springer et al. (2003) contend that sequential declines occurred in North Pacific populations of harbor and fur seals, Steller sea lions, and sea otters. They hypothesize that these were due to increased predation by killer whales, when industrial whaling’s removal of large whales as a supposed primary food source precipitated a prey switch. Using a regional approach, we reexamined whale catch data, killer whale predation observations, and the current biomass and trends of potential prey, and found little support for the prey-switching hypothesis. Large whale biomass in the Bering Sea did not decline as much as suggested by Springer et al., and much of the reduction occurred 50–100 yr ago, well before the declines of pinnipeds and sea otters began; thus, the need to switch prey starting in the 1970s is doubtful. With the sole exception that the sea otter decline followed the decline of pinnipeds, the reported declines were not in fact sequential. Given this, it is unlikely that a sequential megafaunal collapse from whales to sea otters occurred. The spatial and temporal patterns of pinniped and sea otter population trends are more complex than Springer et al. suggest, and are often inconsistent with their hypothesis. Populations remained stable or increased in many areas, despite extensive historical whaling and high killer whale abundance. Furthermore, observed killer whale predation has largely involved pinnipeds and small cetaceans; there is little evidence that large whales were ever a major prey item in high latitudes. Small cetaceans (ignored by Springer et al.) were likely abundant throughout the period. Overall, we suggest that the Springer et al. hypothesis represents a misleading and simplistic view of events and trophic relationships within this complex marine ecosystem.

Demography and population viability of polar bears in the Gulf of Boothia, Nunavut

We estimated demographic parameters and harvest risks for polar bears (Ursus maritimus) inhabiting the Gulf of Boothia, Nunavut, from 1976 to 2000. We computed survival and abundance from capture–recapture and recovery data (630 marks) using a Burnham joint live–dead model implemented in program MARK. Annual mean total survival (including harvest) was 0.889 ± 0.179 ( x ± 1 SE) for cubs, 0.883 ± 0.087 for subadults (ages 1–4), 0.919 ± 0.044 for adult females, and 0.917 ± 0.041 for adult males. Abundance in the last 3 yr of study was 1,592 ± 361 bears. Mean size of newborn litters was 1.648 ± 0.098 cubs. By age 7, 0.97 ± 0.30 of available females were producing litters. Harvest averaged 38.4 ± 4.2 bears/year in the last 5 yr of study; however, the 2002–2007 kill averaged 56.4 bears/yr. We used a harvested Population Viability Analysis (PVA) to examine impacts of increasing rates of harvest. We estimated the current population growth rate, λH, to be 1.025 ± 0.032. Although this suggests the population is growing, progressive environmental changes may require more frequent population inventory studies to maintain the same levels of harvest risk.

Predator and Heterospecific Stimuli Alter Behavior in Cattle

Wild and domestic ungulates modify their behavior in the presence of olfactory and visual cues of predators but investigations have not exposed a domestic species to a series of cues representing various predators and other ungulate herbivores.We used wolf (Canis lupus), mountain lion (Puma concolor), and mule deer (Odocoileus hemionus) stimuli (olfactory and visual), and a control (no stimuli) to experimentally test for differences in behavior of cattle (Bos taurus) raised in Arizona. We measured (1) vigilance, (2) foraging rates, (3) giving up density (GUD) of high quality foods and (4) time spent in high quality forage locations in response to location of stimuli treatments. In general, we found a consistent pattern in that wolf and deer treatments caused disparate results in all 4 response variables. Wolf stimuli significantly increased cattle vigilance and decreased cattle foraging rates; conversely, deer stimuli significantly increased cattle foraging rate and increased cattle use of high quality forage areas containing stimuli. Mountain lion stimuli did not significantly impact any of the 4 response variables. Our findings suggest that domestic herbivores react to predatory stimuli, can differentiate between stimuli representing two predatory species, and suggest that cattle may reduce antipredatory behaviour when near heterospecifics.

Invasive Litter, Not an Invasive Insectivore, Determines Invertebrate Communities in Hawaiian Forests

In Hawaii, invasive plants have the ability to alter litter-based food chains because they often have litter traits that differ from native species. Additionally, abundant invasive predators, especially those representing new trophic levels, can reduce prey. The relative importance of these two processes on the litter invertebrate community in Hawaii is important, because they could affect the large number of endemic and endangered invertebrates. We determined the relative importance of litter resources, represented by leaf litter of two trees, an invasive nitrogen-fixer, Falcataria moluccana, and a native tree, Metrosideros polymorpha, and predation of an invasive terrestrial frog, Eleutherodactylus coqui, on leaf litter invertebrate abundance and composition. Principle component analysis revealed that F. moluccana litter creates an invertebrate community that greatly differs from that found in M. polymorpha litter. We found that F. moluccana increased the abundance of non-native fragmenters (Amphipoda and Isopoda) by 400% and non-native predaceous ants (Hymenoptera: Formicidae) by 200%. E. coqui had less effect on the litter invertebrate community; it reduced microbivores by 40% in F. moluccana and non-native ants by 30% across litter types. E. coqui stomach contents were similar in abundance and composition in both litter treatments, despite dramatic differences in the invertebrate community. Additionally, our results suggest that invertebrate community differences between litter types did not cascade to influence E. coqui growth or survivorship. In conclusion, it appears that an invasive nitrogen-fixing tree species has a greater influence on litter invertebrate community abundance and composition than the invasive predator, E. coqui.

Contributions of demography and dispersal parameters to the spatial spread of a stage-structured insect invasion

Stage-structured models that integrate demography and dispersal can be used to identify points in the life cycle with large effects on rates of population spatial spread, information that is vital in the development of containment strategies for invasive species. Current challenges in the application of these tools include: (1) accounting for large uncertainty in model parameters, which may violate assumptions of ‘‘local’’ perturbation metrics such as sensitivities and elasticities, and (2) forecasting not only asymptotic rates of spatial spread, as is usually done, but also transient spatial dynamics in the early stages of invasion. We developed an invasion model for the Diaprepes root weevil (DRW; Diaprepes abbreviatus [Coleoptera: Curculionidae]), a generalist herbivore that has invaded citrus-growing regions of the United States. We synthesized data on DRW demography and dispersal and generated predictions for asymptotic and transient peak invasion speeds, accounting for parameter uncertainty. We quantified the contributions of each parameter toward invasion speed using a ‘‘global’’ perturbation analysis, and we contrasted parameter contributions during the transient and asymptotic phases. We found that the asymptotic invasion speed was 0.02–0.028 km/week, although the transient peak invasion speed (0.03– 0.045 km/week) was significantly greater. Both asymptotic and transient invasions speeds were most responsive to weevil dispersal distances. However, demographic parameters that had large effects on asymptotic speed (e.g., survival of early-instar larvae) had little effect on transient speed. Comparison of the global analysis with lower-level elasticities indicated that local perturbation analysis would have generated unreliable predictions for the responsiveness of invasion speed to underlying parameters. Observed range expansion in southern Florida (1992–2006) was significantly lower than the invasion speed predicted by the model. Possible causes of this mismatch include overestimation of dispersal distances, demographic rates, and spatiotemporal variation in parameter values. This study demonstrates that, when parameter uncertainty is large, as is often the case, global perturbation analyses are needed to identify which points in the life cycle should be targets of management. Our results also suggest that effective strategies for reducing spread during the asymptotic phase may have little effect during the transient phase. Includes Appendix.

HOME RANGES AND MOVEMENTS OF COYOTES IN THE NORTHERN CHIHUAHUAN DESERT

The coyote (Canis latrans) is among the most studied animals in North America. Because of its adaptability and success as a predator, the coyote has flourished and is still expanding its range. Coyotes can now be found throughout most of North America and south into Central America (Voight and Berg 1987). Studies in recent years have been extensive to understand the interrelationships of prey and coyotes (Shelton and Klindt 1974, Beckoff and Wells 1981), as well as demographic relationships (Davis et al. 1975, Knowlton and Stoddart 1978, Mitchell 1979, Bowen 1981) and feeding strategies (Todd and Keith 1976, Andelt et al. 1987, MacCracken and Hansen 1987, Gese et al. 1988a). With the advance of radio telemetry, researchers have investigated lifestyle characteristics spatially with home ranges or temporally with movements in relation to habitat requirements. Researchers have studied home ranges of coyotes in various regions of the United States (Livaitis and Shaw 1980, Andelt 1981, Springer 1982, Pyrah 1984, Gese et al. 1988a) and Canada (Bowen 1982). Some studies of home range were separated by season (Ozoga and Harger 1966) or relation to nearby food sources (Danner and Smith 1980). Home range analysis in relation to social interactions of coyotes has been either neglected, overlooked, or avoided. Gese et al. (1988a) recognized a transient class of coyote by home range size. Coyote social systems are very complex and can vary by season or locality in addition to some reports of group or pack systems (Hamlin and Schweitzer 1979, Beckoff and Wells 1981, Bowen 1981, Gese et al. 1988b). Coyotes maintain communication with conspecifics through vocal and olfactory signals (Lehner 1987, Bowen and McTaggert Cowan 1980). Social interactions may be by far the most complex and least understood aspect related to coyote ecology. Coyote movements can be related to many factors including food, water, cover, and social interactions. Movements in relation to food sources are well documented (Fitch 1948, Todd and Keith 1976, Danner and Smith 1980) although reports on movements in relation to water have not been reported, probably because of limited research in desert situations. There has been some mention of coyotes' movements in relation to cover (Wells and Beckoff 1982). The objectives of this study were to delineate annual and seasonal home ranges, movements, and habitat use of coyotes in the northern Chihuahuan desert.