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A consumer-resource approach to the density-dependent population dynamics of mutualism

Like predation and competition, mutualism is now recognized as a consumer resource (C-R) interaction, including, in particular, bi-directional (e.g., coral, plant- mycorrhizae) and uni-directional (e.g., ant-plant defense, plant-pollinator) C-R mutualisms. Here, we develop general theory for the density-dependent population dynamics of mutualism based on the C-R mechanism of interspecific interaction. To test the influence of C-R interactions on the dynamics and stability of bi- and uni-directional C-R mutualisms, we developed simple models that link consumer functional response of one mutualistic species with the resources supplied by another. Phase-plane analyses show that the ecological dynamics of C-R mutualisms are stable in general. Most transient behavior leads to an equilibrium of mutualistic coexistence, at which both species densities are greater than in the absence of interactions. However, due to the basic nature of C-R interactions, certain density-dependent conditions can lead to C-R dynamics characteristic of predator-prey interactions, in which one species overexploits and causes the other to go extinct. Consistent with empirical phenomena, these results suggest that the C-R interaction can provide a broad mechanism for understanding density-dependent population dynamics of mutualism. By unifying predation, competition, and mutualism under the common ecological framework of consumer-resource theory, we may also gain a better understanding of the universal features of interspecific interactions in general.

Ecology↗

The distance that contaminated aquatic subsidies extend into lake riparian zones

Consumption of emergent aquatic insects by terrestrial invertebrates is a poorly resolved, but potentially important, mechanism of contaminant flux across ecosystem borders leading to contaminant exposure in terrestrial invertivores. We characterized the spatial extent and magnitude of contaminant transfer from aquatic sediments to terrestrial invertebrate predators by examining riparian araneid spiders, terrestrial insects, and emergent aquatic insects for stable isotopes and polychlorinated biphenyls (PCBs, sum of 141 congeners) at Lake Hartwell, (Clemson, South Carolina, USA). PCB concentrations in aquatic insects were orders of magnitude higher than in terrestrial insects. Aquatic insect consumption by spiders (as indicated by δ 13 C and δ 15 N), PCB concentrations in spiders, and aquatic prey availability were greatest at the shoreline and declined inland, while terrestrial prey availability was invariant with distance. These patterns indicate PCB transfer to spiders through consumption of emergent aquatic insects extending to a distance of ∼5 m inland. Measurable, but much lower, PCBs were present in insect predators dominated by social wasps up to 30 m inland. These results illustrate the importance of emergent insects as vectors of contaminant transfer from lake sediments to riparian food webs, and that spiders are key predators in this process.

South Carolina↗

Evaluating potential conservation conflicts between two listed species: Sea otters and black abalone

Population consequences of endangered species interacting as predators and prey have been considered theoretically and legally, but rarely investigated in the field. We examined relationships between spatially variable populations of a predator, the California sea otter, Enhydra lutris nereis , and a prey species, the black abalone, Haliotis cracherodii . Both species are federally listed under the Endangered Species Act and co-occur along the coast of California. We compared the local abundance and habitat distribution of black abalone at 12 sites with varying densities of sea otters. All of the populations of abalone we examined were in the geographic area currently unaffected by withering disease, which has decimated populations south of the study area. Surprisingly, our findings indicate that sea otter density is positively associated with increased black abalone density. The presence of sea otters also correlated with a shift in black abalone to habitat conferring greater refuge, which could decrease illegal human harvest. These results highlight the need for a multi-species approach to conservation management of the two species, and demonstrate the importance of using field-collected data rather than simple trophic assumptions to understand relationships between jointly vulnerable predator and prey populations.

California↗

Diet shift of lentic dragonfly larvae in response to reduced terrestrial prey subsidies

Inputs of terrestrial plant detritus and nutrients play an important role in aquatic food webs, but the importance of terrestrial prey inputs in determining aquatic predator distribution and abundance has been appreciated only recently. I examined the numerical, biomass, and diet responses of a common predator, dragonfly larvae, to experimental reduction of terrestrial arthropod input into ponds. I distributed paired enclosures ( n = 7), one with a screen between the land and water (reduced subsidy) and one without a screen (ambient subsidy), near the shoreline of 2 small fishless ponds and sampled each month during the growing season in the southern Appalachian Mountains, Virginia (USA). Screens between water and land reduced the number of terrestrial arthropods that fell into screened enclosures relative to the number that fell into unscreened enclosures and open reference plots by 36%. The δ 13 C isotopic signatures of dragonfly larvae shifted towards those of aquatic prey in reduced-subsidy enclosures, a result suggesting that dragonflies consumed fewer terrestrial prey when fewer were available (ambient subsidy: 30%, reduced subsidy: 19% of diet). Overall abundance and biomass of dragonfly larvae did not change in response to reduced terrestrial arthropod inputs, despite the fact that enclosures permitted immigration/emigration. These results suggest that terrestrial arthropods can provide resources to aquatic predators in lentic systems, but that their effects on abundance and distribution might be subtle and confounded by in situ factors.

Journal of the North American Benthological Societ↗

Changing numbers of spawning cutthroat trout in tributary streams of Yellowstone Lake and estimates of grizzly bears visiting streams from DNA

Spawning Yellowstone cutthroat trout ( Oncorhynchus clarki ) provide a source of highly digestible energy for grizzly bears ( Ursus arctos ) that visit tributary streams to Yellowstone Lake during the spring and early summer. During 1985–87, research documented grizzly bears fishing on 61% of the 124 tributary streams to the lake. Using track measurements, it was estimated that a minimum of 44 grizzly bears fished those streams annually. During 1994, non-native lake trout ( Salvelinus namaycush ) were discovered in Yellowstone Lake. Lake trout are efficient predators and have the potential to reduce the native cutthroat population and negatively impact terrestrial predators that use cutthroat trout as a food resource. In 1997, we began sampling a subset of streams ( n = 25) from areas of Yellowstone Lake surveyed during the previous study to determine if changes in spawner numbers or bear use had occurred. Comparisons of peak numbers and duration suggested a considerable decline between study periods in streams in the West Thumb area of the lake. The apparent decline may be due to predation by lake trout. Indices of bear use also declined on West Thumb area streams. We used DNA from hair collected near spawning streams to estimate the minimum number of bears visiting the vicinity of spawning streams. Seventy-four individual bears were identified from 429 hair samples. The annual number of individuals detected ranged from 15 in 1997 to 33 in 2000. Seventy percent of genotypes identified were represented by more than 1 sample, but only 31% of bears were documented more than 1 year of the study. Sixty-two (84%) bears were only documented in 1 segment of the lake, whereas 12 (16%) were found in 2–3 lake segments. Twenty-seven bears were identified from hair collected at multiple streams. One bear was identified on 6 streams in 2 segments of the lake and during 3 years of the study. We used encounter histories derived from DNA and the Jolly-Seber procedure in Program MARK to produce annual estimates of grizzly bears visiting streams. Approximately 68 grizzly bears visited the vicinity of cutthroat trout spawning streams annually. Thus, approximately 14–21% of grizzly bears in the Greater Yellowstone Ecosystem (GYE) may have used this threatened food resource annually. Yellowstone National Park (YNP) is attempting to control the lake trout population in Yellowstone Lake; our results underscore the importance of that effort to grizzly bears.

Wyoming↗

Does presence of permanent fresh water affect recruitment in prairie-nesting dabbling ducks?

In the Prairie Pothole Region (PPR) of North Dakota, USA, American mink ( Mustela vison ) are a major predator of ducklings. Mink populations plummet during severe droughts, but some mink survive where permanent fresh water is available. In 1992–1993, we evaluated whether development of a permanent water body, the 125-km McClusky Canal (MC), had affected survival of gadwall ( Anas strepera ) and mallard ( A. platyrhynchos ) broods and ducklings in surrounding wetland complexes. Twelve of 25 radiomarked gadwall and mallard hens experienced total brood loss, and 148 of 199 radiomarked ducklings from 58 broods died by day 30. Gadwall broods ( n = 18 radiomarked hens) survived to 30 days at a lower rate (0.52) than predicted for similar areas in the region with limited permanent fresh water (0.85; P = 0.009). Observed ( n = 162 radiomarked ducklings from 48 broods) survival rates also were lower than predicted for gadwall ducklings 0–7 days old (0.42 vs. 0.60; P < 0.001) and 8–30 days old (0.41 vs. 0.80; P < 0.001). We attempted to include mallards in models constructed to predict brood and duckling survival rates in the Koenig Study Area (KSA), but data were too sparse. Rates of survival to 30 days for gadwall and mallard ducklings declined from an estimated 0.83 and 0.68 in 1976–1981 (Lokemoen et al. 1990), when the MC was first filling with water, to 0.36 and 0.31 (adjusted for radiotransmitter effects) in 1992–1993 after the MC had become a permanent freshwater body. Estimated gadwall recruitment rate (females fledged per hen) during 1992–1993 was 0.5, <50% of the estimated recruitment rate in 1976–1981. Of 130 radiomarked ducklings (both species) for which we determined cause of death, 114 mortalities were attributed to predation; at least 65% of 62 deaths in which the predator type could be discerned were caused by mink. Environmental planners and waterfowl managers should be aware of potential risks to waterfowl production from development of permanent freshwater bodies in prairie pothole landscapes and may wish to refine duck productivity models to consider negative effects of permanent water on duckling survival.

North Dakota↗

Nest survival of clay-colored and vesper sparrows in relation to woodland edge in mixed-grass prairies

The quantity and quality of northern mixed-grass prairie continues to decline because of conversion to agriculture, invasion of woody and exotic plants, and disruption of important ecological processes that shape grasslands. Declines in grassland bird populations in North Dakota, USA, have coincided with these largely anthropogenic alterations to prairie habitat. In grasslands of north-central and northwestern North Dakota, woody plants have increased due primarily to fire suppression, extirpation of bison (Bos bison), and widescale planting of tree shelter belts. In northern grasslands, effects of woody vegetation on survival of grassland birds are poorly understood, and conclusions are based mainly on studies conducted outside the region. We examined nest survival of clay-colored sparrows (Spizella pallida) and vesper sparrows (Pooecetes gramineus) relative to the distance nests were located from aspen (Populus tremuloides,) woodland edges and relative to other habitat features near the nest. Clay-colored and vesper sparrow nest survival was higher for nests located near woodland edges, nests with greater cover of Kentucky bluegrass (Poa pratensis), and nests more concealed by vegetation. Vesper sparrow nest survival increased as the percent cover of tall shrubs near the nest increased. Based on video-camera data, the 13-lined ground squirrel (Spermophilus tridecemlineatus,) was the most common predator of sparrow eggs and young. Thirteen-lined ground squirrels were more common far from woodland edges than near, and this pattern may, in part, explain clay-colored and vesper sparrow nest survival in relation to woodland edges. In contrast to our results, studies conducted in other grassland systems generally report lower nest survival for grassland birds nesting near trees and shrubs. This disparity in results demonstrates the need to identify specific nest predators and their distributions with respect to important habitat features because these data can be important in explaining-and perhaps predicting-patterns of nest predation.

Journal of Wildlife Management↗

Survival rates, mortality causes, and habitats of Pennsylvania white-tailed deer fawns

Estimates of survival and cause-specific mortality of white-tailed deer ( Odocoileus virginianus ) fawns are important to population management. We quantified cause-specific mortality, survival rates, and habitat characteristics related to fawn survival in a forested landscape and an agricultural landscape in central Pennsylvania. We captured and radiocollared neonatal (<3 weeks) fawns in 2000–2001 and monitored fawns from capture until death, transmitter failure or collar release, or the end of the study. We estimated survivorship functions and assessed influence on fawn survival of road density, habitat edge density, habitat patch diversity, and proportion of herbaceous habitat. We captured 110 fawns in the agricultural landscape and 108 fawns in the forested landscape. At 9 weeks after capture, fawn survival was 72.4% (95% Cl = 63.3–80.0%) in the agricultural landscape and 57.2% (95% Cl = 47.5–66.3%) in the forested landscape. Thirty-four-week survival was 52.9% (95% Cl = 42.7–62.8%) in the agricultural landscape and 37.9% (95% Cl = 27.7–49.3%) in the forested landscape. We detected no relationship between fawn survival and road density, percent herbaceous cover, habitat edge density, or habitat patch diversity (all P >0.05). Predation accounted for 46.2% (95% Cl = 37.6–56.7%) of 106 mortalities through 34 weeks. We attributed 32.7% (95% Cl = 21.9–48.6%) and 36.7% (95% Cl = 25.5–52.9%) of 49 predation events to black bears ( Ursus americanus ) and coyotes ( Canis latrans ), respectively. Natural causes, excluding predation, accounted for 27.4% (95% Cl = 20.1–37.3) of mortalities. Fawn survival in Pennsylvania was comparable to reported survival in forested and agricultural regions in northern portions of the white-tailed deer range. We have no evidence to suggest that the fawn survival rates we observed were preventing population growth. Because white-tailed deer are habitat generalists, home-range-scale habitat characteristics may be unrelated to fawn survival; therefore, future studies should consider landscape-related characteristics on fawn survival.

Pennsylvania↗

Ranking Alaska moose nutrition: Signals to begin liberal antlerless harvests

We focused on describing low nutritional status in an increasing moose ( Alces alces gigas ) population with reduced predation in Game Management Unit (GMU) 20A near Fairbanks, Alaska, USA. A skeptical public disallowed liberal antlerless harvests of this moose population until we provided convincing data on low nutritional status. We ranked nutritional status in 15 Alaska moose populations (in boreal forests and coastal tundra) based on multiyear twinning rates. Data on age-of-first-reproduction and parturition rates provided a ranking consistent with twinning rates in the 6 areas where comparative data were available. Also, short-yearling mass provided a ranking consistent with twinning rates in 5 of the 6 areas where data were available. Data from 5 areas implied an inverse relationship between twinning rate and browse removal rate. Only in GMU 20A did nutritional indices reach low levels where justification for halting population growth was apparent, which supports prior findings that nutrition is a minor factor limiting most Alaska moose populations compared to predation. With predator reductions, the GMU 20A moose population increased from 1976 until liberal antlerless harvests in 2004. During 1997–2005, GMU 20A moose exhibited the lowest nutritional status reported to date for wild, noninsular, North American populations, including 1) delayed reproduction until moose reached 36 months of age and the lowest parturition rate among 36-month-old moose (29%, n = 147); 2) the lowest average multiyear twinning rates from late-May aerial surveys ( x̄ = 7%, SE = 0.9%, n = 9 yr, range = 3–10%) and delayed twinning until moose reached 60 months of age; 3) the lowest average mass of female short-yearlings in Alaska ( x̄ = 155 ± 1.6 [SE] kg in the Tanana Flats subpopulation, up to 58 kg below average masses found elsewhere); and 4) high removal (42%) of current annual browse biomass compared to 9–26% elsewhere in boreal forests. When average multiyear twinning rates in GMU 20A (sampled during 1960–2005) declined to <10% in the mid- to late 1990s, we began encouraging liberal antlerless harvests, but only conservative annual harvests of 61–76 antlerless moose were achieved during 1996–2001. Using data in the context of our broader ranking system, we convinced skeptical citizen advisory committees to allow liberal antlerless harvests of 600–690 moose in 2004 and 2005, with the objective of halting population growth of the 16,000–17,000 moose; total harvests were 7–8% of total prehunt numbers. The resulting liberal antlerless harvests served to protect the moose population's health and habitat and to fulfill a mandate for elevated yield. Liberal antlerless harvests appear justified to halt population growth when multiyear twinning rates average ≤10% and ≥1 of the following signals substantiate low nutritional status: <50% of 36-month-old moose are parturient, average multiyear short-yearling mass is <175 kg, or >35% of annual browse biomass is removed by moose.

Journal of Wildlife Management↗

Factors influencing nesting success of king eiders on northern Alaska's Coastal Plain

King eider (Somateria spectabilis) populations have declined markedly in recent decades for unknown reasons. Nest survival is one component of recruitment, and a female's chance of reproductive success increases with her ability to choose an appropriate nesting strategy. We estimated variation in daily nest survival of king eiders at 2 sites, Teshekpuk and Kuparuk, Alaska, USA, 2002-2005. We evaluated both a priori and exploratory competing models of nest survival that considered importance of nest concealment, seclusion, and incubation constancy as strategies to avoid 2 primary egg predators, avian (Larus spp., Stercorarius spp., and Corvus corax) and fox (Alopex lagopus). We used generalized nonlinear techniques to examine factors affecting nest survival rates and information-theoretic approaches to select among competing models. Estimated nest survival, accounting for a nest visitation effect, varied considerably across sites and years (0.21-0.57); however, given our small sample size, much of this variation maybe attributable to sampling variation (??process = 0.007, 95% CI: 0.003-0.070). Nest survival was higher at Kuparuk than Teshekpuk in all years; however, due to the correlative nature of our data, we cannot determine the underlying causes with any certainty. We found mixed support for the concealed breeding strategy, females derived no benefit from nesting in areas with more willow (Salix spp.; measure of concealment) except that the observer effect diminished as willow cover increased. We suggest these patterns are due to conflicting predation pressures. Nest survival was not higher on islands (measure of seclusion) or with increased incubation constancy but was higher post-fox removal, indicating that predator control on breeding grounds could be a viable management option. Nest survival was negatively affected by our nest visitations, most likely by exposing the nest to avian scavengers. We recommend precautions be taken to limit the effects of nest visits in future studies and to consider them as a possible negative bias in estimated nest survival. Future models of the impacts of development within the breeding grounds of king eider should consider the influence of humans in the vicinity of nests.

Journal of Wildlife Management↗

Factors influencing bird foraging preferences among conspecific fruit trees

The rates at which birds visit fruiting individuals of Allophylus edulis (Sapindaceae) differ substantially among trees. Such avian feeding preferences are well-known, but usually involve fruits and trees of different species. Factors controlling avian preferences for particular trees in a population of conspecifics are generally undocumented. To address this issue, I attempted to correlate rates at which individual birds and species fed in trees of Allophylus with 27 fruit or plant characteristics. Birds that swallow fruits whole were considered separately from those that feed in other ways. Plant characters were selected on the basis of their potential influence on feeding efficiency or predation risk, assuming that birds would select feeding trees so as to maximize the net rate of energy or nutrient intake and to minimize predation. Correlations were found between feeding visits by some groups of birds and percent water in the pulp, milligrams of mineral ash in the pulp, and crop size. No character was correlated with feeding visits by all groups of birds in both years of the study. The correlations with water and mineral ash are unexplained and may be artifacts. The correlation with crop size may represent a tactic to minimize predation.

Condor↗

Acadian flycatcher nest placement: Does placement influence reproductive success?

We located 511 Acadian Flycatcher (Empidonax virescens) nests in bottomland hardwood forest of eastern Arkansas. Microhabitat characteristics were measured and their relationship with nest success evaluated. Fifty-two percent of all nesting attempts resulted in predation. Attributes of nest placement were similar between successful and unsuccessful nests, although successful nests were placed higher. Similarly, nonparasitized nests were typically higher than parasitized nests. Nests initiated late in the breeding season were placed in larger trees with higher canopy bases resulting in increased vegetation around the nest. Fifteen different tree species were used for nesting. Acadian Flycatchers chose nest trees in a nonrandom fashion, selecting Nuttall oak (Quercus nuttallii) and possumhaw (Ilex decidua) in greater proportions than their availability. However, there was no relationship between tree species used for nesting and nest success. Nest height was positively correlated with concealment at the nest site, supporting the predator-avoidance theory. No other attribute of nest placement differentiated successful nest sites, suggesting that nest predation is likely a function of random events in space and time.

Condor↗

Relative effects of plumage coloration and vegetation density on nest success

Many passerine species are highly dichromatic with brightly-colored males and cryptically-colored females. Bright plumage in males is commonly thought to arise as a result of sexual selection by females such that males with bright coloration possess high fitness. However, bright plumage potentially could expose males to increased predation risk. Consistent with this idea, males of many highly dichromatic passerine species do not incubate. I tested whether brightly-colored males avoid incubation to reduce the probability of visual predators locating their nest. This hypothesis predicts greater hatching success for clutches incubated by cryptically-colored individuals than by brightly-colored individuals. The Northern Cardinal (Cardinalis cardinalis) is a common dichromatic species that breeds throughout the eastern U.S. I placed two button-quail (Turnix st).) eggs in each of 203 simulated cardinal nests. Dull brown cardboard, simulating a female cardinal, was placed over about half of all clutches. Bright red cardboard, simulating a male cardinal, was placed over the other clutches. Nest success was highest for well-concealed nests (87%) and lowest for nests in open habitat (54%). Nests containing red cardboard did not have significantly lower success than nests with brown cardboard, nor did I detect a significant color X vegetation-density interaction. My analysis may have had insufficient power to detect an effect of color on nest success; alternatively, brightly-colored males that do not incubate may achieve benefits unrelated to predation risk.

Condor↗

Life-history strategies of ungulates

This Special Feature resulted from a symposium on life-history strategies of ungulates presented at the 78th Annual Meeting of the American Society of Mammalogists in Blacks-burg, Virginia, in June 1998. The presentations at the symposium represented only a vignette of the wide variety of life-history strategies that exists among ungulates. The four papers that follow include treatises on birth-site selection of moose ( Alces alces ), sex-ratio correlates with dimorphism and risk of predation, optimal foraging relative to risk of predation, and the role of density dependence in shaping life-history traits of ungulates. A theme of risk of predation in shaping life-history traits is common to three of four papers.

Journal of Mammalogy↗

Sea otters and kelp forests in Alaska: Generality and variation in a community ecological paradigm

Multiscale patterns of spatial and temporal variation in density and population structure were used to evaluate the generality of a three—trophic—level cascade among sea otters (Enhydra lutris), invertebrate herbivores, and macroalgae in Alaska. The paradigm holds that where sea otters occur herbivores are rare and plants are abundant, whereas when sea otters are absent herbivores are relatively common and plants are rare. Spatial patterns were based on 20 randomly placed quadrats at 153 randomly selected sites distributed among five locations with and four locations without sea otters. Both sea urchin and kelp abundance differed significantly among locations with vs. without sea otters in the Aleutian Islands and southeast Alaska. There was little (Aleutian Islands) or no (southeast Alaska) overlap between sites with and without sea otters, in plots of kelp density against urchin biomass. Despite intersite variation in the abundance of kelps and herbivores, these analyses demonstrate that sea otter predation has a predictable and broadly generalizable influence on the structure of Alaskan kelp forests. The percent cover of algal turf and suspension feeder assemblages also differed significantly (although less dramatically) between locations with and without sea otters. Temporal variation in community structure was assessed over periods of from 3 to 15 yr at sites in the Aleutian Islands and southeast Alaska where sea otters were 1) continuously present, 2) continuously absent, or 3) becoming reestablished because of natural range expansion. Kelp and sea urchin abundance remained largely unchanged at most sites where sea otters were continuously present or absent, the one exception being at Torch Bay (southeast Alaska), where kelp abundance varied significantly through time and urchin abundance varied significantly among sites because of episodic and patchy disturbances. In contrast, kelp and sea urchin abundances changed significantly, and in the expected directions, at sites that were being recolonized by sea otters. Sea urchin biomass declined by 50% in the Aleutian Islands and by nearly 100% in southeast Alaska following the spread of sea otters into previously unoccupied habitats. In response to these different rates and magnitudes of urchin reduction by sea otter predation, increases in kelp abundance were abrupt and highly significant in southeast Alaska but much smaller and slower over similar time periods in the Aleutian Islands. The different kelp colonization rates between southeast Alaska and the Aleutian Islands appear to be caused by large—scale differences in echinoid recruitment coupled with size—selective predation by sea otters for larger urchins. The length of urchin jaws (correlated with test diameter, r 2 = 0.968) in sea otter scats indicates that sea urchins <15—20 mm test diameter are rarely eaten by foraging sea otters. Sea urchin populations in the Aleutian Islands included high densities of small individuals (<20 mm test diameter) at all sites and during all years sampled, whereas in southeast Alaska similarly sized urchins were absent from most populations during most years. Small (<30—35 mm test diameter) tetracycline—marked urchins in the Aleutian Islands grew at a maximum rate of °10 mm/yr; thus the population must have significant recruitment annually, or at least every several years. In contrast, echinoid recruitment in southeast Alaska was more episodic, with many years to perhaps decades separating significant events. Our findings help explain regional differences in recovery rates of kelp forests following recolonization by sea otters.

Alaska↗

Effects of introducing foxes and raccoons on herring gull colonies

Red foxes ( Vulpes fulva ) and raccoons ( Procyon lotor ) released at colonies of herring gulls ( Larus argentatus ) on islands off the Massachusetts coast effectively eliminated the production of young gulls. Annual predator introductions for 2-4 years caused major reductions in colony size and occasionally total abandonment of the island as a colony site. Observations of the experimental islands for 2 years after cessation of predator introductions showed slow repopulation of the islands and lower breeding success than on control islands. The size of the regional population was reduced largely because of the movements of gulls off the experimental islands. The introduced predators are, in most cases, difficult to maintain on the islands; this restricts their utility in population management.

Massachusetts↗

Mathematics and mallard management

Waterfowl managers can effectively use simple population models to aid in making management decisions. We present a basic model of the change in population size as related to survival and recruitment. A management technique designed to increase survival of mallards (Anas platyrhynchos) by limiting harvest on the Chippewa National Forest, Minnesota, is used to illustrate the application of models in decision making. The analysis suggests that the management technique would be of limited effectiveness. In a 2nd example, the change in mallard population in central North Dakota is related to implementing programs to create dense nesting cover with or without supplementary predator control. The analysis suggests that large tracts of land would be required to achieve a hypothetical management objective of increasing harvest by 50% while maintaining a stable population. Less land would be required if predator reduction were used in combination with cover management, but questions about effectiveness and ecological implications of large scale predator reduction remain unresolved. The use of models as a guide to planning research responsive to the needs of management is illustrated.

Journal of Wildlife Management↗

Reproductive performance of female Alaskan caribou

We examined the reproductive performance of female caribou ( Rangifer tarandus granti ) in relation to age, physical condition, and reproductive experience for 9 consecutive years (1987-95) at Denali National Park, Alaska, during a period of wide variation in winter snowfall. Caribou in Denali differed from other cervid populations where reproductive performance has been investigated, because they occur at low densities (≥0.3/km 2 ) and experience high losses of young to predation. Females first gave birth at 2-6 years old; 56% of these females were 3 years old. Average annual natality rates increased from 27% for 2-year-olds to 100% for 7-year-olds, remained high for 7-13-year-olds (98%), and then declined for females ≥14 years old. Females ≥2 years old that failed to reproduce were primarily sexually immature (76%). Reproductive pauses of sexually mature females occurred predominantly in young (3-6 yr old) and old (≥14 yr old) females. Natality increased with body mass for 10-month-old females weighed 6 months prior to the autumn breeding season (P = 0.007), and for females >1 year old and weighed during autumn (late Sep-early Nov; P = 0.003). Natality for 2-, 3-, 4-, and 6-year-olds declined with increasing late-winter snowfall (Feb-May; P ≤ 0.039) during the winter prior to breeding. In most years, a high percentage of sexually mature females reproduced, and lactation status at the time of breeding did not influence productivity the following year. However, following particularly high snowfall during February-September 1992, productivity was reduced in 1993 for cows successfully rearing calves to autumn the previous year. High losses of calves to predators in 1992 may have increased productivity in 1993. Losses of young-of-the-year to predation prior to the annual breeding season can be an important influence on subsequent productivity for ungulate populations where productivity varies with lactation status of females at the time of breeding.

Journal of Wildlife Management↗