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Contributions of vital rates to growth of a protected population of American black bears

Analyses of large, long-lived animals suggest that adult survival generally has the potential to contribute more than reproduction to population growth rate (λ), but because survival varies little, high variability in reproduction can have a greater influence. This pattern has been documented for several species of large mammals, but few studies have evaluated such contributions of vital rates to λ for American black bears ( Ursus americanus ). We used variance-based perturbation analyses (life table response experiments, LTRE) and analytical sensitivity and elasticity analyses to examine the actual and potential contributions of variation of vital rates to variation in growth rate (λ) of a population of black bears inhabiting the Pisgah Bear Sanctuary in the southern Appalachian Mountains of North Carolina, using a 22-year dataset. We found that recruitment varied more than other vital rates; LTRE analyses conducted over several time intervals thus indicated that recruitment generally contributed at least as much as juvenile and adult survival to observed variation in λ, even though the latter 2 vital rates had the greater potential to affect λ. Our findings are consistent with predictions from studies on polar bears ( U. maritimus ) and grizzly bears ( U. arctos ), but contrast with the few existing studies on black bears in ways that suggest levels of protection from human-caused mortality might explain whether adult survival or recruitment contribute most to variation in λ for this species. We hypothesize that λ is most strongly influenced by recruitment in protected populations where adult survival is relatively high and constant, whereas adult survival will most influence λ for unprotected populations.

Ursus↗

Detecting occurrence and timing of grizzly bear parturition based on anomalies in accelerometer data

Documenting natality of brown (grizzly) bears ( Ursus arctos ) is an important component of many population monitoring and research programs. Brown bears give birth during hibernation, so observation of litters is generally not feasible until females exit dens, and detection of litters can be compromised by poor observability. Using triaxial accelerometer activity data stored on radiocollars, we developed a technique that predicts births by the presence of brief upsurges in activity likely associated with postnatal maternal behaviors. We developed presence criteria with a training sample of known parturient females ( n = 22) to detect anomalies within time-series of daily motion-counts (i.e., activity readings >0 measured at 10-min intervals) during 25 December–7 March. To test performance, we applied criteria to a blinded sample of activity data obtained from female grizzly bears (captured and collared during 2012–2022) in 4 populations in interior North America ( n = 295). We assigned predicted status and compared assignments to reproductive status at first visual observation. The true positive rate was 91% ( n = 47 females observed with cubs of the year) and the false-positive rate was 17% ( n = 65 females observed with older offspring). Births were predicted for 50% of females with unknown reproductive status ( n = 48), 21% of females observed without offspring ( n = 114), and 10% of subadult females considered too young to reproduce ( n = 21). Dates of predicted births varied from 27 December to 28 February. Our anomaly detection technique was successful at estimating parturition events, and despite some error, indicated that a number of litters were born but not observed, presumably due to mortality in or shortly after den emergence. This technique provides an additional tool for supplementing visual observations for natality estimation and population modeling, provided that potential biases stemming from increased detection rates are considered.

Alaska, British Columbia, Idaho, Montana↗

Nesting ecology of boreal forest birds following a massive outbreak of spruce beetles

We studied breeding dark-eyed juncos ( Junco hyemalis ), yellow-rumped warblers ( Dendroica coronata ), and spruce-nesting birds from 1997 to 1998 among forests with different levels of spruce ( Picea spp.) mortality following an outbreak of spruce beetles ( Dendroctonus rufipennis ) in Alaska, USA. We identified species using live and beetle-killed spruce for nest sites and monitored nests to determine how the outbreak influenced avian habitat selection and reproduction. We tested predictions that 1) nesting success of ground-nesting juncos would increase with spruce mortality due to proliferation of understory vegetation available to conceal nests from predators, 2) nesting success of canopy-nesting warblers would decrease with spruce mortality due to fewer live spruce in which to conceal nests, and 3) both species would alter nest-site selection in response to disturbance. Juncos did not benefit from changes in understory vegetation; nesting success in highly disturbed stands (46%) was comparable to that in undisturbed habitats throughout their range. In stands with low spruce mortality, nesting success of juncos was low (5%) and corresponded with high densities of red squirrels ( Tamiasciurus hudsonicus ). Yellow-rumped warblers nested exclusively in spruce, but success did not vary with spruce mortality. As disturbance increased, nesting warblers switched from selecting forest patches with high densities of live white spruce ( Picea glauca ) to patches with beetle-killed spruce. Warblers also placed nests in large-diameter live or beetle-killed spruce, depending on which was more abundant in the stand, with no differences in nesting success. Five of the 12 other species of spruce-nesting birds also used beetle-killed spruce as nest sites. Because beetle-killed spruce can remain standing for >50 years, even highly disturbed stands provide an important breeding resource for boreal forest birds. We recommend that boreal forest managers preserve uncut blocks of infested forest within managed forest landscapes and practice partial harvest of beetle-killed spruce rather than commercial clear-cutting of infested stands in order to sustain breeding bird populations until natural reforestation occurs. Because breeding densities do not always reflect fitness, assessing impacts of a massive natural disturbance should include measuring impacts of changes in vegetation on both reproductive success and predator–prey dynamics.

Journal of Wildlife Management↗

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↗

The disappearance of Guam's wildlife: New insights for herpetology, evolutionary ecology, and conservation

The wealth of data generated from intensive study of the brown tree snake as a result of the need to control introduced populations of this pest species allow several important conclusions. First, that the snakes on Guam are extraordinary in terms of their absolute abundance and in terms of their ability to exploit a broad prey base. Our data suggest an exceptionally high reproductive success on Guam for a snake with an otherwise unnoteworthy reproductive capability and life history (i.e. small clutch size, typical ontogenetic shift from small heterothermic prey to larger homeotherms). Especially important was the snakes versatility in taking advantage of extremely common prey on islands; population expansion was slow but survival was maximal, ultimately leading to high population levels. The brown tree snake shares many attributes with other snakes that could cause similar biodiversity crises in a wide variety of contexts in which they lack coevolutionary histories (especially formerly snake-free island environments). As opposed to their relatively poor history as over-water dispersers, snakes may be especially problematic as travelers in increasing ship and air traffic between widely separated geographic regions of the world.

BioScience↗

Organochlorine residues, eggshell thickness, and nest success in barn owls from the Chesapeake Bay

Eggs of barn owls ( Tyto alba ) were collected from 18 nests in offshore duck blinds on the Maryland side of the lower Potomac River estuary in 1972 and 1973 and analyzed for organochlorine residues. DDE was found in 100% of the clutches, PCBs in 89%, and dieldrin in 78%. Eggshell thickness was inversely correlated with concentrations of DDE, DDD, and dieldrin residues. Six of the 18 clutches had mean DDE residues above 5 ppm, and eggshell thickness in these six clutches was significantly less (P<0.001) than in the other 12 clutches. The owls produced 1.7 young per active nest in 1973. This rate is slightly below the reproductive rate needed to maintain a stable population. An estimated 15% of the population carried concentrations of organochlorine residues that may have been detrimental to their reproduction. Passerine birds, taken extensively as food by a small proportion of the population, are believed to have been the source of elevated concentrations of organochlorines in these barn owls.

Maryland, Virginia↗

Variation in egg size of the northern pintail

Egg size is an important determinant of reproductive investment by birds. For many species, total investment in a clutch is limited by the size of stored reserves (Ankney and MacInnes 1978, Esler and Grand 1994a). Egg size determines the unit by which these stored reserves are partitioned. Individual females in most species of waterfowl show a high repeatability for egg size, implying that individual either cannot, or do not, alter their egg size in response to varying environmental conditions (batt and Prince 1979, Duncan 1987, Laurila and Hario 1988, Lessells et al 1989, Flint and Sedinger 1992). Thus differences in egg size appear to represent different reproductive strategies among individuals. Fitness can be measured by the number of offspring an individual contributes to a population. Egg size may be related to fitness in some species fo waterfowl as young from larger eggs are better able to survive extreme conditions (Ankney 1980, Thomas and Brown 1988). Birds laying larger clutches are almost always more fit as they fledge more young (Lessells 1986, Rockwell et al 1987, Flint 1993). These fitness patterns create the potential for a trade-off between clutch size and egg size where females laying large clutches of small eggs have the same fitness as females laying smaller clutches of large eggs. The fact that Northern Pintails ( Anas acuta ) utilize stored reserves (Mann and Sedinger 1993, esler and Grand 1994a) and have a high repeatability for egg size (i.e. egg size is fixed) (Duncan 1987), makes them candidates to engage in clutch size=egg size trade-offs (Rowher 1988, Rowher and Eisenhauer 1989). An inverse relationship between egg size and clutch size would be indicative of a phenotypic trade-off among these fitness components. Our goal in this study was to describe egg size variation in Northern Pintails (hereafter pintails) with regard to female age, body size, clutch size, year, initiation date, and nesting attempt. We compare our results to those from other populations of nesting pintails and discuss whether phenotypic clutch size-egg size tradeoffs exist for pintails.

Alaska↗

Trends and effects of organochlorine residues on Oregon and Nevada wading birds, 1979-83

The incidence of DDT (parent compound) declined significantly in the eggs of Black-crowned Night-Herons (Nycticorax nycticorax), White-faced Ibis (Plegadis chihi), and Black-necked Stilts (Himantopus mexicanus), and showed a downward trend in Snowy Egrets (Egretta thula). Mean DDE residues (breakdown product of DDT) in eggs declined significantly in Black-crowned Night-Herons and showed a downward trend in Black-necked Stilts and Snowy Egrets; however, no trend was apparent for White-faced Ibis. Some night-herons, egrets, and ibis laid thin-shelled eggs that cracked; eggshell thickness was negatively correlated with DDE residues in the egg contents. Ibis were the most sensitive species to DDE- induced shell thinning. Generally, reduced reproductive success for ibis started at lower DDE egg residues (3 ppm) than for either Snowy Egrets (5 ppm) or night-herons (8 ppm). Egg residue profiles, band recoveries, biotelemetry studies, and residues in fish and other prey items implicate the southwestern United States (a wintering area) as an important source of DDT-DDE that caused the most serious reproductive problem encountered during the study (in night-herons nesting at Ruby Lake, Nevada). Available evidence indicates that the three other species, together with night- herons from Oregon and Idaho, winter in Latin America.

Idaho, Nevada, Oregon↗

Environmental contaminants, human disturbance and nesting of double-crested cormorants in northwestern Washington

Double-crested Cormorants (Phalacrocorax auritus) in extreme northwestern Washington produced few young (0.27/occupied nest) in 1984; the clutch size was generally small and eggs, if laid at all, were laid later than usual. Residues (geometric means, wet weight) of DDE (0.58 and 0.59 ppm) in eggs from Colville Island and Protection Island were lower than from other locations in the Pacific Northwest, while PCBs (2.19 and 1.37 ppm) were similar to those at most locations. Both contaminants in 1984 were below levels associated with reproductive problems. Eggs also contained concentrations of mercury (0.26 and 0.27 ppm) and selenium (0.31 and 0.28 ppm) below levels associated with reproductive problems. The distribution of nesting colonies in the study area changed dramatically since 1984. The cormorants were most likely responding to increased human disturbance in the San Juan Islands, coupled to additional protection and reduced human activity on Protection and Smith Islands. This presumably led to the abandonment of all nesting islands in the San Juans. The nesting population in the study area in 1988 (all on Protection and Smith Islands) was the highest recorded.

Colonial Waterbirds↗

Monitoring arctic habitat and goose production by satellite imagery

Spacecraft imagery, especially from the National Atmospheric and Oceanic Administration's Improved TIROS (Television Infra-Red Observational Satellite) Operational Satellites, permits timely evaluations of snow and ice conditions encountered by arctic nesting geese. Imagery from the TIROS satellite for 5 wide]y scattered locations in arctic North America was obtained for three 3-day intervals in June 1973 and 1974. These pictures were used to expand fragmentary habitat data available from ground observations. Late disappearance of snow and ice may prevent or retard nesting effort and reproductive success. Our immediate aim is to recognize years of catastrophic or very good production; however, supporting information from ground studies, LANDSAT imagery, analyses of banding data, and studies of age ratios in popu]ations and harvests eventua]]y may permit assessment of relative reproductive success. Satellite images from both Multi-Spectral Scanner and Very High Resolution Radiomder sensors were first used as management tools for consideration in setting the U.S. and Canadian regulations for the 1975-76 goose hunting season. Preliminary age ratio data from field observations and harvests substantiated the prediction of good production during the 1975 breeding season.

Journal of Wildlife Management↗

Feeding ecology of breeding gadwalls on saline wetlands

The feeding ecology of breeding gadwalls ( Anas strepera ) from saline wetlands in North Dakota was examined in relation to sex, pair mates, reproductive status, food availability, and wetland type during the spring and summer of 1971 and 1972. Esophagi of males and females contained 40.4 and 48.2 percent animal food, respectively, between 17 April and 25 August. Animal foods consumed by paired females varied with reproductive condition and were independent of their mates. Invertebrates increased from 47.7 i?? 17.4 percent in the diet during prelaying to 72.0 i?? 18.4 percent during laying and declined to 46.3 i?? 30.0 percent during postlaying. Aquatic insects dominated the diet during egg-laying and were selected disproportionately relative to their availability. Esophageal contents indicated that diversity of plant and animal foods in the diet varied inversely with specific conductance. Major factors influencing food selection of the breeding birds are discussed as interactions among their physiological status, their anatomical and behavioral characteristics, and the abundance and behavior of food organisms as influenced by chemical and physical features of the environment. The data suggested that these interrelated ecological factors act simultaneously to control the phenology of events and determine the foods utilized.

North Dakota↗

Foods of breeding pintails in North Dakota

Food habits of breeding pintails ( Anas acuta ) were studied relative to sex, land use, and reproductive condition during the spring and summer of 1969, 1970, and 1971 in eastern North Dakota. Hens and drakes, respectively, consumed 79.2 percent and 30.0 percent animal matter on nontilled wetlands and consumed 16.6 percent and 1.1 percent animal matter on tilled wetlands. Aquatic dipterans (primarily larval forms), snails, fairy shrimp, and earthworms accounted for 71 percent of the diet of hens on nontilled wetlands, while barnyard grass ( Echinochloa crusgalli ) seeds formed 71 percent of the diet of hens on tilled wetlands. Cereal grain seeds formed 84 percent of the diet of 10 hens feeding on cropland. The diet of hens was influenced by reproductive status. Animal foods were predominant during the laying period (77.1 percent) but were less important in the postlaying diet (28.9 percent). Invertebrates formed 83.9 percent of the diet of renesting hens, 61.0 percent were dipteran larvae and snails. High consumption of animal foods during egg formation presumably is related to invertebrates being superior to plants in providing certain nutrients required for production of viable eggs. Research findings suggest that food requirements of prairie-nesting pintails can be met most effectively by providing pairs access to shallow, nontilled wetland habitat subject to periodic drawdowns.

North Dakota↗

Effects of heptachlor- and lindane-treated seed on Canada geese

A study of Canada geese (B. canadensis) was conducted in the Pacific Northwest from 1978 through 1981. Lowered reproductive success, mortality of adults, and a population decline of resident western Canada geese (B. c. moffitti) at the Umatilla National Wildlife Refuge (Umatilla NWR) Oregon and Washington [USA] were associated with the use of heptachlor-treated wheat. Analyses of eggs and tissues from dead geese provided strong evidence that heptachlor was responsible for the mortality and decreased production. Residues of heptachlor epoxide (HE) in brains of B. c. moffitti found dead in 1978 and 1979 equalled or exceeded the lethal hazard zone of 8-9 .mu.g/g in experimental passerine birds; HE residues of > 10 .mu.g/g in sample eggs were associated with low nest success. The breeding population of Canada geese at Umatilla decreased from 129 pairs in 1974 to about 100 pairs in 1979. The heptachlor problem was restricted to the Umatilla area; HE residues were low in tissues and eggs collected at other sites in Oregon, Washington and Idaho. In Sept., 1979, the use of heptachlor-treated seed was banned in a 1700-km2 area that encompassed both sides of the Columbia River near Umatilla. Lindane was substituted for heptachlor in the restricted area in 1979, and its use was extended to much of the Columbia Basin in 1981. Concurrently, reproductive success of geese increased, mortality decreased, and the nesting population increased to 170 pairs by 1983. There was no evidence for either biomagnification of lindance residues from treated seed to goose tissues or eggs or for induction of adverse effects by this compound.

Oregon, Washington↗

Survival of radiomarked canvasback ducklings in northwestern Minnesota

Duckling survival, an important factor affecting annual recruitment, has not been determined adequately for canvasbacks ( Aythya valisineria ). We investigated the magnitude, timing, and causes of mortality of canvasback ducklings from hatch to fledging at the Agassiz National Wildlife Refuge (NWR) in northwestern Minnesota during 1987-90. During the 4 years, 217 day-old ducklings were radiomarked and released in 52 broods. Another 141 ducklings were radiomarked at ≥ to 4 weeks of age. Survival was estimated with the Kaplan-Meier nonparametric estimator and the Weibull parametric model. Most mortalities occurred within 10 days after hatch. Total brood loss occurred in 18 (85%) of 52 broods released. The primary sources of mortality were predation principally by mink ( Mustela vison ), and exposure to precipitation and cold temperature. For combined years, females had lower survival than males ( P =0.03). If the disparate survival between sexes of canvasbacks observed in this study is representative of canvasbacks in their breeding range, this phenomenon contributes to reduced reproductive potential and the male-biased sex ratio of the species.

Minnesota↗

A model of northern pintail productivity and population growth rate

Our objective was to synthesize individual components of reproductive ecology into a single estimate of productivity and to assess the relative effects of survival and productivity on population dynamics. We used information on nesting ecology, renesting potential, and duckling survival of northern pintails ( Anas acuta ) collected on the Yukon-Kuskokwim Delta (Y-K Delta), Alaska, 1991-95, to model the number of ducklings produced under a range of nest success and duckling survival probabilities. Using average values of 25% nest success, 11% duckling survival, and 56% renesting probability from our study population, we calculated that all young in our population were produced by 13% of the breeding females, and that early-nesting females produced more young than later-nesting females. Further, we calculated, on average, that each female produced only 0.16 young females/nesting season. We combined these results with estimates of first-year and adult survival to examine the growth rate (X) of the population and the relative contributions of these demographic parameters to that growth rate. Contrary to aerial survey data, the population projection model suggests our study population is declining rapidly (X = 0.6969). The relative effects on population growth rate were 0.1175 for reproductive success, 0.1175 for first-year survival, and 0.8825 for adult survival. Adult survival had the greatest influence on X for our population, and this conclusion was robust over a range of survival and productivity estimates. Given published estimates of annual survival for adult females (61%), our model suggested nest success and duckling survival need to increase to approximately 40% to achieve population stability. We discuss reasons for the apparent discrepancy in population trends between our model and aerial surveys in terms of bias in productivity and survival estimates.

Alaska↗

Food use and nutrition of black ducks nesting in Maine

Based on 32 adult black ducks ( Anas rubripes ) collected during the nesting seasons of 1974-76, the proportion of macro invertebrates (as aggregate [average] percent of dry weight) in the diet of males, egg-laying females, and postlaying females was 60, 75, and 55%, respectivey. Sample sizes were small, and the differences associated with sex and reproductive condition were not signficant. Molluscs, Isopods, ephemeropteran and odonate nymphs, and coleopteran, trichopteran, and dipteran larvae contributed 74% of the dry weight and 64% of the gross energy ingested. Data from proximate analyses of 9 invertebrate and 9 plant foods were combined with food habits data to estimate the nutrients available to breeding hens. We concluded that females with access to an adequate amount of natural food including invertebrates and the seeds and tubers of aquatic plants would obtain sufficient minerals, protein, and energy for reproduction. Food quality does not appear to limit the density of black ducks nesting in Maine.

Maine↗

Selenium and boron in aquatic birds from central California

Subsurface agricultural drainwater used for marsh management has resulted in trace element contamination of aquatic bird food chains in central California. Consequently, we collected breeding and wintering aquatic birds from the Grassland Water District (GWD) of California during 1985-88 to measure selenium (Se) and boron (B) contamination resulting from use of such drainage water for wetland management. During the breeding and wintering periods, livers of birds from the North and South areas of the Grasslands contained concentrations of Se and B that have been associated with reproductive impairment. Birds from the South Grasslands, which had received more undiluted drainage water, were more contaminated than those from the North Grasslands. Birds had higher (P < 0.001) levels of Se and B at the end of the 1985-86 wintering period than at the beginning, indicating that the Grasslands was the major source of contamination. Concentrations of Se decreased from 1985 through 1988, after freshwater was substituted for irrigation drainage water during autumn 1985. B concentrations in wintering birds, except for American coots (Fulica americana), declined to background levels, while concentrations in breeding birds remained slightly elevated. However, after 3 years of freshwater management of the Grasslands, liver Se levels in some breeding and wintering birds still were above concentrations associated with impaired reproduction in laboratory and field studies. In areas with high potential for leaching of Se and B from agricultural land, irrigation drainage water should not be used for wetland management.

California↗

Effects of tillage practices and carbofuran exposure on small mammals

We compared population estimates, body mass, movement, and blood chemistry of small mammals between conventionally tilled and no-till cornfields in Maryland and Pennsylvania to evaluate the effects of tillage practices and carbofuran exposure on small mammals. Estimates suggest that populations of white-footed mice ( Peromyscus leucopus ) were not significantly related (P > 0.05) to tillage practice or to the presence of winter rye cover crops. Late summer differences in animal mass and movement, and in the proportion of reproductively inactive animals imply that characteristics of individuals and reproductive activity can be affected by tillage practices and winter cover crops. The presence of granular carbofuran had no apparent effect on populations, body mass, or movement in conventionally tilled or no-till fields. Similarly, neither red blood cell acetylcholinesterase, hematocrit, nor liver function were affected by carbofuran use.

Maryland, Pennsylvania↗