Artificial intelligence for automated detection of large mammals creates path to upscale drone surveys.

Imagery from drones is becoming common in wildlife research and management, but processing data efficiently remains a challenge. We developed a methodology for training a convolutional neural network model on large-scale mosaic imagery to detect and count caribou (Rangifer tarandus), compare model performance with an experienced observer and a group of naïve observers, and discuss the use of aerial imagery and automated methods for large mammal surveys. Combining images taken at 75 m and 120 m above ground level, a faster region-based convolutional neural network (Faster-RCNN) model was trained in using annotated imagery with the labels: "adult caribou", "calf caribou", and "ghost caribou" (animals moving between images, producing blurring individuals during the photogrammetry processing). Accuracy, precision, and recall of the model were 80%, 90%, and 88%, respectively. Detections between the model and experienced observer were highly correlated (Pearson: 0.96-0.99, P value < 0.05). The model was generally more effective in detecting adults, calves, and ghosts than naïve observers at both altitudes. We also discuss the need to improve consistency of observers' annotations if manual review will be used to train models accurately. Generalization of automated methods for large mammal detections will be necessary for large-scale studies with diverse platforms, airspace restrictions, and sensor capabilities.

Kin grouping is insufficient to explain the inclusive fitness gains of conspecific brood parasitism in the common eider.

Conspecific brood parasitism allows females to exploit other females' nests and enhance their reproductive output. Here, we test a recent theoretical model of how host females gain inclusive fitness from brood parasitism. High levels of relatedness between host and parasitizer can be maintained either by: (a) kin recognizing and parasitizing each other as a form of cooperative breeding or (b) natal philopatry and nest site fidelity facilitating the formation of kin groups, thereby increasing the probability of parasitism between relatives nesting in close proximity. To address these two hypotheses we genotyped feathers and hatch membranes of common eiders (Somateria mollissima) from western Hudson Bay, Canada, using a noninvasive sampling methodology. We found that most instances of brood parasitism do result in inclusive fitness gains. Furthermore, females with failed nests moved an average of 492 m from their previous year's nest site, while successful females only moved an average of 13 m. Therefore, we observed host-parasite relatedness can occur at levels higher than would be expected by chance even in the absence of kin grouping, suggesting that closely related females nesting near one another is not essential to maintain high host-parasitizer relatedness. In addition, kin grouping is only a transient phenomenon that cannot occur every year due to the propensity for females of failed nests to nest farther away from their nest site in subsequent years than females with successful nests, which provides support for kin recognition as a more likely mechanism to maintain high host-parasitizer relatedness over time.

A comparison of drone imagery and ground-based methods for estimating the extent of habitat destruction by lesser snow geese (Anser caerulescens caerulescens) in La Pérouse Bay.

Lesser snow goose (Anser caerulescens caerulescens) populations have dramatically altered vegetation communities through increased foraging pressure. In remote regions, regular habitat assessments are logistically challenging and time consuming. Drones are increasingly being used by ecologists to conduct habitat assessments, but reliance on georeferenced data as ground truth may not always be feasible. We estimated goose habitat degradation using photointerpretation of drone imagery and compared estimates to those made with ground-based linear transects. In July 2016, we surveyed five study plots in La Pérouse Bay, Manitoba, to evaluate the effectiveness of a fixed-wing drone with simple Red Green Blue (RGB) imagery for evaluating habitat degradation by snow geese. Ground-based land cover data was collected and grouped into barren, shrub, or non-shrub categories. We compared estimates between ground-based transects and those made from unsupervised classification of drone imagery collected at altitudes of 75, 100, and 120 m above ground level (ground sampling distances of 2.4, 3.2, and 3.8 cm respectively). We found large time savings during the data collection step of drone surveys, but these savings were ultimately lost during imagery processing. Based on photointerpretation, overall accuracy of drone imagery was generally high (88.8% to 92.0%) and Kappa coefficients were similar to previously published habitat assessments from drone imagery. Mixed model estimates indicated 75m drone imagery overestimated barren (F2,182 = 100.03, P < 0.0001) and shrub classes (F2,182 = 160.16, P < 0.0001) compared to ground estimates. Inconspicuous graminoid and forb species (non-shrubs) were difficult to detect from drone imagery and were underestimated compared to ground-based transects (F2,182 = 843.77, P < 0.0001). Our findings corroborate previous findings, and that simple RGB imagery is useful for evaluating broad scale goose damage, and may play an important role in measuring habitat destruction by geese and other agents of environmental change.

Shifting Vital Rate Correlations Alter Predicted Population Responses to Increasingly Variable Environments.

Time series of vital rates are often used to construct "environment-blind" stochastic population projections and calculate the elasticity of population growth to increased temporal variance in vital rates. Here, we show that the utility of this widely used demographic tool is greatly limited by shifts in vital rate correlations that occur as environmental drivers become increasingly variable. The direction and magnitude of these shifts are unpredictable without environmentally explicit models. Shifting vital rate correlations had the largest fitness effects on life histories with short to medium generation times, potentially hampering comparative analyses based on elasticities to vital rate variance for a wide range of species. Shifts in vital rate correlations are likely ubiquitous in increasingly variable environments, and further research should empirically evaluate the life histories for which detailed mechanistic relationships between vital rates and environmental drivers are required for making reliable predictions versus those for which summarized demographic data are sufficient.

Reproductive success of a keystone herbivore is more variable and responsive to climate in habitats with lower resource diversity.

The effects of climate on wild populations are often channelled through species interactions. Population responses to climate variation can therefore differ across habitats, owing to variation in the biotic community. Theory predicts that consumer demography should be less variable and less responsive to climate in habitats with greater resource diversity. We tested these predictions using a long-term study of breeding lesser snow geese along the western coast of Hudson Bay, Manitoba, Canada. Reproductive success was measured in 22 years from 114 locations, in either coastal or inland habitat types. We used Bayesian analysis to estimate the response of reproductive success to climate in each habitat type, along with residual variation not explained by climate. We then quantified gosling diet composition in each habitat type to test the prediction that reproductive success would be less variable and more responsive to climate in habitats with lower resource diversity. Reproductive success responded positively to seasonal warmness, but this response was much stronger in inland habitats than in coastal habitats. Site- and year-level random effects were also three to five times more variable in inland habitats. Simultaneously, land cover diversity and gosling diet diversity were lower in inland habitats. Our study illustrates that spatial variation in resource diversity (and thus, species interactions) can have important effects on consumer responses to climate. In this system, climate change is expected to disproportionately increase the reproductive success of snow geese in vast inland habitats, potentially counteracting management efforts to reduce the abundance of this keystone herbivore.

Evaluating behavioral responses of nesting lesser snow geese to unmanned aircraft surveys.

Unmanned aircraft systems (UAS) are relatively new technologies gaining popularity among wildlife biologists. As with any new tool in wildlife science, operating protocols must be developed through rigorous protocol testing. Few studies have been conducted that quantify the impacts UAS may have on unhabituated individuals in the wild using standard aerial survey protocols. We evaluated impacts of unmanned surveys by measuring UAS-induced behavioral responses during the nesting phase of lesser snow geese (Anser caerulescens caerulescens) in Wapusk National Park, Manitoba, Canada. We conducted surveys with a fixed-wing Trimble UX5 and monitored behavioral changes via discreet surveillance cameras at 25 nests. Days with UAS surveys resulted in decreased resting and increased nest maintenance, low scanning, high scanning, head-cocking and off-nest behaviors when compared to days without UAS surveys. In the group of birds flown over, head-cocking for overhead vigilance was rarely seen prior to launch or after landing (mean estimates 0.03% and 0.02%, respectively) but increased to 0.56% of the time when the aircraft was flying overhead suggesting that birds were able to detect the aircraft during flight. Neither UAS survey altitude nor launch distance alone in this study was strong predictors of nesting behaviors, although our flight altitudes (≥75 m above ground level) were much higher than previously published behavioral studies. Synthesis and applications: The diversity of UAS models makes generalizations on behavioral impacts difficult, and we caution that researchers should design UAS studies with knowledge that some minimal disturbance is likely to occur. We recommend flight designs take potential behavioral impacts into account by increasing survey altitude where data quality requirements permit. Such flight designs should consider a priori knowledge of focal species' behavioral characteristics. Research is needed to determine whether any such disturbance is a result of visual or auditory stimuli.

Increased variance in temperature and lag effects alter phenological responses to rapid warming in a subarctic plant community.

Summer temperature on the Cape Churchill Peninsula (Manitoba, Canada) has increased rapidly over the past 75 years, and flowering phenology of the plant community is advanced in years with warmer temperatures (higher cumulative growing degree days). Despite this, there has been no overall shift in flowering phenology over this period. However, climate change has also resulted in increased interannual variation in temperature; if relationships between phenology and temperature are not linear, an increase in temperature variance may interact with an increase in the mean to alter how community phenology changes over time. In our system, the relationship between phenology and temperature was log-linear, resulting in a steeper slope at the cold end of the temperature spectrum than at the warm end. Because below-average temperatures had a greater impact on phenology than above-average temperatures, the long-term advance in phenology was reduced. In addition, flowering phenology in a given year was delayed if summer temperatures were high the previous year or 2 years earlier (lag effects), further reducing the expected advance over time. Phenology of early-flowering plants was negatively affected only by temperatures in the previous year, and that of late-flowering plants primarily by temperatures 2 years earlier. Subarctic plants develop leaf primordia one or more years prior to flowering (preformation); these results suggest that temperature affects the development of flower primordia during this preformation period. Together, increased variance in temperature and lag effects interacted with a changing mean to reduce the expected phenological advance by 94%, a magnitude large enough to account for our inability to detect a significant advance over time. We conclude that changes in temperature variability and lag effects can alter trends in plant responses to a warming climate and that predictions for changes in plant phenology under future warming scenarios should incorporate such effects.

Costs of locomotion in polar bears: when do the costs outweigh the benefits of chasing down terrestrial prey?

Trade-offs between locomotory costs and foraging gains are key elements in determining constraints on predator-prey interactions. One intriguing example involves polar bears pursuing snow geese on land. As climate change forces polar bears to spend more time ashore, they may need to expend more energy to obtain land-based food. Given that polar bears are inefficient at terrestrial locomotion, any extra energy expended to pursue prey could negatively impact survival. However, polar bears have been regularly observed engaging in long pursuits of geese and other land animals, and the energetic worth of such behaviour has been repeatedly questioned. We use data-driven energetic models to examine how energy expenditures vary across polar bear mass and speed. For the first time, we show that polar bears in the 125-235 kg size range can profitably pursue geese, especially at slower speeds. We caution, however, that heat build-up may be the ultimate limiting factor in terrestrial chases, especially for larger bears, and this limit would be reached more quickly with warmer environmental temperatures.

The Energetic Value of Land-Based Foods in Western Hudson Bay and Their Potential to Alleviate Energy Deficits of Starving Adult Male Polar Bears.

Climate change is predicted to expand the ice-free season in western Hudson Bay and when it grows to 180 days, 28-48% of adult male polar bears are projected to starve unless nutritional deficits can be offset by foods consumed on land. We updated a dynamic energy budget model developed by Molnar et al. to allow influx of additional energy from novel terrestrial foods (lesser snow geese, eggs, caribou) that polar bears currently consume as part of a mixed diet while on land. We calculated the units of each prey, alone and in combination, needed to alleviate these lethal energy deficits under conditions of resting or limited movement (2 km d-1) prior to starvation. We further considered the total energy available from each sex and age class of each animal prey over the period they would overlap land-bound polar bears and calculated the maximum number of starving adult males that could be sustained on each food during the ice-free season. Our results suggest that the net energy from land-based food, after subtracting costs of limited movement to obtain it, could eliminate all projected nutritional deficits of starving adult male polar bears and likely other demographic groups as well. The hunting tactics employed, success rates as well as behavior and abundance of each prey will determine the realized energetic values for individual polar bears. Although climate change may cause a phenological mismatch between polar bears and their historical ice-based prey, it may simultaneously yield a new match with certain land-based foods. If polar bears can transition their foraging behavior to effectively exploit these resources, predictions for starvation-related mortality may be overestimated for western Hudson Bay. We also discuss potential complications with stable-carbon isotope studies to evaluate utilization of land-based foods by polar bears including metabolic effects of capture-related stress and consuming a mixed diet.

Effects of exploitation on an overabundant species: the lesser snow goose predicament.

Invasive and overabundant species are an increasing threat to biodiversity and ecosystem functioning world-wide. As such, large amounts of money are spent each year on attempts to control them. These efforts can, however, be thwarted if exploitation is compensated demographically or if populations simply become too numerous for management to elicit an effective and rapid functional response. We examined the influence of these mechanisms on cause-specific mortality in lesser snow geese using multistate capture-reencounter methods. The abundance and destructive foraging behaviours of snow geese have created a trophic cascade that reduces (sub-) Arctic plant, insect and avian biodiversity, bestowing them the status of 'overabundant'. Historically, juvenile snow geese suffered from density-related degradation of their saltmarsh brood-rearing habitat. This allowed harvest mortality to be partially compensated by non-harvest mortality (process correlation between mortality sources: ρ = -0.47; 90% BCI: -0.72 to -0.04). Snow goose family groups eventually responded to their own degradation of habitat by dispersing to non-degraded areas. This relaxed the pressure of density dependence on juvenile birds, but without this mechanism for compensation, harvest began to have an additive effect on overall mortality (ρ = 0.60; 90% BCI: -0.06 to 0.81). In adults, harvest had an additive effect on overall mortality throughout the 42-year study (ρ = 0.24; 90% BCI: -0.59 to 0.67). With the aim of controlling overabundant snow geese, the Conservation Order amendment to the International Migratory Bird Treaty was implemented in February of 1999 to allow for harvest regulations that had not been allowed since the early 1900s (e.g. a spring harvest season, high or unlimited bag limits and use of electronic calls and unplugged shotguns). Although harvest mortality momentarily increased following these actions, the increasing abundance of snow geese has since induced a state of satiation in harvest that has driven harvest rates below the long-term average. More aggressive actions will thus be needed to halt the growth and spread of the devastating trophic cascade that snow geese have triggered. Our approach to investigating the impacts of population control efforts on cause-specific mortality will help guide more effective management of invasive and overabundant species world-wide.

Dietary composition and spatial patterns of polar bear foraging on land in western Hudson Bay.

BACKGROUND: Flexible foraging strategies, such as prey switching, omnivory and food mixing, are key to surviving in a labile and changing environment. Polar bears (Ursus maritimus) in western Hudson Bay are versatile predators that use all of these strategies as they seasonally exploit resources across trophic levels. Climate warming is reducing availability of their ice habitat, especially in spring when polar bears gain most of their annual fat reserves by consuming seal pups before coming ashore in summer. How polar bears combine these flexible foraging strategies to obtain and utilize terrestrial food will become increasingly important in compensating for energy deficits from lost seal hunting opportunities. We evaluated patterns in the composition of foods in scat to characterize the foraging behaviors that underpin the diet mixing and omnivory observed in polar bears on land in western Hudson Bay. Specifically, we measured diet richness, proportions of plant and animal foods, patterns in co-occurrence of foods, spatial composition and an index of temporal composition. RESULTS: Scats contained between 1 and 6 foods, with an average of 2.11 (SE = 0.04). Most scats (84.9%) contained at least one type of plant, but animals (35.4% of scats) and both plants and animals occurring together (34.4% of scats) were also common. Certain foods, such as Lyme grass seed heads (Leymus arenarius), berries and marine algae, were consumed in relatively higher proportions, sometimes to the exclusion of others, both where and when they occurred most abundantly. The predominance of localized vegetation in scats suggests little movement among habitat types between feeding sessions. Unlike the case for plants, no spatial patterns were found for animal remains, likely due the animals' more vagile and ubiquitous distribution. CONCLUSIONS: Our results suggest that polar bears are foraging opportunistically in a manner consistent with maximizing intake while minimizing energy expenditure associated with movement. The frequent mixing of plant-based carbohydrate and animal-based protein could suggest use of a strategy that other Ursids employ to maximize weight gain. Further, consuming high rates of certain vegetation and land-based animals that may yield immediate energetic gains could, instead, provide other benefits such as fulfilling vitamin/mineral requirements, diluting toxins and assessing new foods for potential switching.

What to eat now? Shifts in polar bear diet during the ice-free season in western Hudson Bay.

Under current climate trends, spring ice breakup in Hudson Bay is advancing rapidly, leaving polar bears (Ursus maritimus) less time to hunt seals during the spring when they accumulate the majority of their annual fat reserves. For this reason, foods that polar bears consume during the ice-free season may become increasingly important in alleviating nutritional stress from lost seal hunting opportunities. Defining how the terrestrial diet might have changed since the onset of rapid climate change is an important step in understanding how polar bears may be reacting to climate change. We characterized the current terrestrial diet of polar bears in western Hudson Bay by evaluating the contents of passively sampled scat and comparing it to a similar study conducted 40 years ago. While the two terrestrial diets broadly overlap, polar bears currently appear to be exploiting increasingly abundant resources such as caribou (Rangifer tarandus) and snow geese (Chen caerulescens caerulescens) and newly available resources such as eggs. This opportunistic shift is similar to the diet mixing strategy common among other Arctic predators and bear species. We discuss whether the observed diet shift is solely a response to a nutritional stress or is an expression of plastic foraging behavior.

Climate change, phenology, and habitat degradation: drivers of gosling body condition and juvenile survival in lesser snow geese.

Nesting migratory geese are among the dominant herbivores in (sub) arctic environments, which have undergone unprecedented increases in temperatures and plant growing days over the last three decades. Within these regions, the Hudson Bay Lowlands are home to an overabundant breeding population of lesser snow geese that has dramatically damaged the ecosystem, with cascading effects at multiple trophic levels. In some areas the overabundance of geese has led to a drastic reduction in available forage. In addition, warming of this region has widened the gap between goose migration timing and plant green-up, and this 'mismatch' between goose and plant phenologies could in turn affect gosling development. The dual effects of climate change and habitat quality on gosling body condition and juvenile survival are not known, but are critical for predicting population growth and related degradation of (sub) arctic ecosystems. To address these issues, we used information on female goslings marked and measured between 1978 and 2005 (4125 individuals). Goslings that developed within and near the traditional center of the breeding colony experienced the effects of long-term habitat degradation: body condition and juvenile survival declined over time. In newly colonized areas, however, we observed the opposite pattern (increase in body condition and juvenile survival). In addition, warmer than average winters and summers resulted in lower gosling body condition and first-year survival. Too few plant 'growing days' in the spring relative to hatch led to similar results. Our assessment indicates that geese are recovering from habitat degradation by moving to newly colonized locales. However, a warmer climate could negatively affect snow goose populations in the long-run, but it will depend on which seasons warm the fastest. These antagonistic mechanisms will require further study to help predict snow goose population dynamics and manage the trophic cascade they induce.

Predators, alternative prey and climate influence annual breeding success of a long-lived sea duck.

1. Perturbations to ecosystems have the potential to directly and indirectly affect species interactions, with subsequent impacts on population dynamics and the vital rates that regulate them. 2. The few long-term studies of common eider breeding ecology indicate that reproductive success is low in most years, interrupted by occasional boom years. However, no study has explicitly examined the drivers of long-term variation in reproductive success. 3. Here, we use encounter history data collected across 41 years to examine the effects of arctic foxes (a terrestrial nest predator), local abundance and spatial distribution of lesser snow geese (an alternative prey source), and spring climate on common eider nest success. 4. Eider nest success declined over the course of the study, but was also highly variable across years. Our results supported the hypothesis that the long-term decline in eider nest success was caused by apparent competition with lesser snow geese, mediated by shared predators. This effect persisted even following a large-scale exodus of nesting geese from the eider colony. Nest success was also lowest in years of low arctic fox index, presumably driven by prey switching in years of low small mammal availability. However, increased snow goose abundance appeared to buffer this effect through prey swamping. The effect of spring climate depended on the stage of the breeding season; cold and wet and warm and dry conditions in early spring were correlated with decreased nest success, whereas warm and wet conditions in late spring increased eider nest success. 5. These results underscore the significance of both trophic interactions and climate in regulating highly variable vital rates, which likely have important consequences for population dynamics and the conservation of long-lived iteroparous species.

Rapid environmental degradation in a subarctic ecosystem influences resource use of a keystone avian herbivore.

1. Environmental degradation can change resource use strategies of animals and thereby affect survival and fitness. Arctic herbivores may be especially susceptible to the effects of such environmental change because their rapid growth rates demand high-quality forage, which may be limited as environmental conditions deteriorate. We studied the consequences of a trophic cascade, driven by Lesser Snow Goose (Chen caerulescens caerulescens) overgrazing on the south-west coast of Hudson Bay, Canada, which has caused tidal marsh (TM) degradation and the reduction in high-quality forage plants, on gosling growth and resource use. 2. We compared resource use and body size of goslings that inhabited tidal and freshwater marsh (FM) to determine how current foraging strategies influence growth and to test the hypothesis that during early growth goslings require and so consume high-quality TM plants, but that during later growth they may switch to foraging in lower-quality FM. 3. To investigate gosling resource use throughout growth, we measured once a week for 28 days the body size of goslings as well as stable isotope ratios (δ(34) S, δ(15) N and δ(13) C) in multiple tissues of goslings that were collected from both TM and nearby FM. We also measured the stable isotope ratios in forage plants sampled along transects and from gosling foreguts. We used an isotope-mixing model to determine the contribution of FM plants to gosling tissues. 4. Contrary to the proposed hypothesis, goslings inhabiting FM or TM primarily consumed FM plants during early growth. Furthermore, goslings that foraged extensively in FM had similar growth rates and grew to a similar size and body mass, as goslings that foraged in the degraded TM. However, goslings that currently inhabit freshwater or TM were significantly smaller than goslings that inhabited TM in the 1980s prior to habitat degradation. 5. Consequences of smaller overall body size include decreased survival and fecundity for arctic-nesting geese. The ability of phenotypically plastic responses to sustain persistence is limited by reaction norms and the extent of environmental change. Current research is assessing whether those limits have been reached in this system.

Inference of population structure and patterns of gene flow in canine heartworm (Dirofilaria immitis).

Understanding the genetic variation within a parasitic species is crucial to implementing successful control programs and preventing the dispersal of drug resistance alleles. We examined the population genetics and structure of canine heartworm (Dirofilaria immitis) by developing a panel of 11 polymorphic microsatellite loci for this abundant parasite. In total, 192 individual nematodes were opportunistically sampled from 9 geographic regions in the United States and Mexico and genotyped. Population genetic analyses indicate the presence of 4 genetic clusters. The canine heartworm samples used in this study were characterized by low heterozygosity, with eastern and central North America experiencing high levels of reciprocal gene flow. Geographic barriers impede the movement of vectors and infected hosts west of the Rocky Mountains and south of the Central Mexican Plateau. This, combined with corridors of contiguous habitat, could influence the spread of drug resistance alleles.

Effects of landscape features on the distribution and sustainability of ungulate hunting in northern Congo.

Understanding the spatial dimensions of hunting and prey population dynamics is important in order to estimate the sustainability of hunting in tropical forests. We investigated how hunting offtake of vertebrates differed in mixed forest and monodominant forest (composed of Gilbertiodendron dewevrei) and over different spatial extents within the hunting catchment around the logging town of Kabo, Congo. In 9 months of recall surveys with hunters, we gathered information on over 1500 hunting trips in which ungulates were 65% of the species killed and 82% of harvested biomass. Hunters supplied information on animals killed and the hunting trip, including the area visited (i.e., hunting zone; 11 separate zones within a 506 km(2) catchment or commonly hunted area). Over 65% of all animals were killed in monodominant forest, which made up 28% of the hunting catchment, and zones with small amounts of monodominant forest were used most frequently by hunters. Given the large offtakes from monodominant forests, we suggest that animal dispersal may be maintaining high, localized harvests in these areas. We believe hunters preferred to hunt in monodominant forest because the understory was accessible and that areas with small amounts of monodominant forest and large amounts of mixed forest were more productive. The variation in hunting pressure we found between and within hunting zones differs from past examinations of spatial variation in hunting offtake, where entire hunting catchments were considered population sinks and areas with low to no hunting (no-take zones) were outside hunting catchments. Future use of no-take zones to manage hunting should incorporate variability in offtake within hunting catchments.

Ugandan Kaposi's sarcoma-associated herpesvirus phylogeny: evidence for cross-ethnic transmission of viral subtypes.

OBJECTIVE: The aim of this study was to test the relationship between Kaposi's sarcoma-associated herpesvirus (KSHV) phylogeny and host ethnicity at the within-country scale. METHODS: KSHV genomic DNA samples were isolated from 31 patients across eleven Ugandan ethnic groups. Amino acid sequences of the ORF-K1 gene were used to construct a neighbor-joining phylogenetic tree. RESULTS: A5 and B1 variants predominated with no evidence of distinct ethnic or geographic distribution. A new K1 subtype (F) was identified in a member of the Bantu Gisu tribe and a new subtype B variant (B3) among members of the Bantu Ganda tribe. CONCLUSIONS: The phylogeny may yet be structured by host ethnicity if members of Ugandan groups have convoluted biological origins, even as they identify with single tribes. An alternative possibility is that KSHV subtype evolution may have preceded major diversification of sub-Saharan Africans into ethnicities as we know them today, with ethnic groups beginning their histories already hosting multiple subtypes. A third alternative is that horizontal transmission of multiple KSHV subtypes may have broken up vertical lineages of the virus passed down within Ugandan populations.

FREQUENCY-DEPENDENT FITNESS CONSEQUENCES OF INTRASPECIFIC NEST PARASITISM IN SNOW GEESE.

The reproductive efficiency, defined as the number of breeding recruits produced per egg laid; of intraspecific nest parasites; of hosts in parasitized nests; and of unparasitized nesting females, was measured for 14 years for lesser snow geese Anser caerulescens caerulescens nesting near Churchill, Manitoba, Canada. Relative efficiencies were 0.71-0.88, 0.91, and 1.0 for eggs of parasites, hosts, and unparasitized birds, respectively. Differences in the hatching probabilities of the three classes of eggs produced the efficiency differences. Parasitic success was limited by the parasites' failure to place more eggs than expected by chance into nests at the appropriate time relative to host incubation. Host nesting success was lower when more than one parasitic egg was added to the clutch. No differences in gosling survival and breeding recruitment probabilities were detected among any categories of goslings. Thus, hatching parasitic young are at no disadvantage relative to parental young, and there is no support for the hypothesis that increased success of host young at later stages of reproduction might offset negative effects at the egg stage. The hatching efficiency of parasitic eggs declined more rapidly than that of parental eggs as the parasitism rate increased. Efficiencies were similar when 3-4% of the eggs laid per year were parasitic, but relative parasitic efficiency was significantly lower when the parasitism rate was 8-9% or more. Using ancillary information and assumptions about the fecundity, viability, and behavioral flexibility of parasitic and parental females, we conclude that intraspecific nest parasitism could compete with nesting as a reproductive strategy in this population. The conditional use of parasitism by a large component of the population in certain years, however, combined with negative-frequency dependent success, limits the potential spread of a purely parasitic strategy in this population.

LIFE HISTORY STUDIES OF THE LESSER SNOW GOOSE (ANSER CAERULESCENS CAERULESCENS). IV. THE SELECTIVE VALUE OF PLUMAGE POLYMORPHISM: NET VIABILITY, THE TIMING OF MATURATION, AND BREEDING PROPENSITY.

Cooch (1961, 1963) suggested that changes in the genotypic composition of snow goose colonies comprising the Hudson Bay-Foxe Basin population could be attributed to selection favoring the blue phenotype. In the preceding paper (Cooke et al., 1985), we examined potential differences in net fecundity between the two phenotypes at La Pérouse Bay in northern Manitoba. No substantial differences in any component of fecundity were detected. In the present paper, we examined potential differences in pre-reproductive and adult viability, age of maturation, and breeding propensity. Again, no differences associated with the plumage polymorphism were detected in any of these life history characteristics. Thus, despite a thorough analysis of the complete life cycle, we were unable to uncover any evidence supporting a selection hypothesis for the persistence and dynamics of this conspicuous polymorphism. An alternate hypothesis based primarily on gene flow and assortative mating appears more plausible.