Sexual Size Dimorphism in Hawks and Owls of North America
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We conducted a meta-analysis to provide a current assessment of the population characteristics of California Spotted Owls (Strix occidentalis occidentalis) resident on four study areas in the Sierra Nevada and one study area in southern California. Our meta-analysis followed rigorous a priori analysis protocols, which we derived through extensive discussion during a week-long analysis workshop. Because there is great interest in the owl?s population status, we used state-of-the-art analytical methods to obtain results as precise as possible. Our meta-analysis included data from five California study areas located on the Lassen National Forest (1990-2000), Eldorado National Forest (1986-2000), Sierra National Forest (1990-2000), Sequoia and Kings Canyon national parks (1990-2000), and San Bernardino National Forest (1987-1998). Four of the five study areas spanned the length of the Sierra Nevada, whereas the fifth study area encompassed the San Bernardino Mountains in southern California. Study areas ranged in size from 343 km2 (Sequoia and Kings Canyon) to 2,200 km (Lassen). All studies were designed to use capture-recapture methods and analysis. We used survival in a meta-analysis because field methods were very similar among studies. However, we did not use reproduction in a meta-analysis because it was not clear if variation among individual study-area protocols used to assess reproductive output of owls would confound results. Thus, we analyzed fecundity only by individual study area. We examined population trend using the reparameterized Jolly-Seber capture-recapture estimator (8t) We did not estimate juvenile survival rates because of estimation problems and potential bias because of juvenile emigration from study areas. We used mark-recapture estimators under an information theoretic framework to assess apparent survival rates of adult owls. The pooled estimate for adult apparent survival for the five study areas was 0.833, which was lower than pooled adult survival rates (0.850) from 15 Northern Spotted Owl (S. o. caurina) studies. Estimates of survival from the best model on the Lassen (N = 0.829, 95% confidence intervals [CI = 0.798 to 0.857), Eldorado (N = 0.815, 95% CI = 0.772 to 0.851), Sierra (N = 0.818, 95% CI = 0.781 to 0.850), and San Bernardino (N = 0.813, 95% CI = 0.782 to 0.841) were not different. However, the Sequoia and Kings Canyon population had a higher survival rate (N = 0.877, 95% CI = 0.842 to 0.905) than the other study areas. Management history and forest structure (e.g. presence of giant sequoia [Sequoiadendron giganteum]) on the Sequoia and Kings Canyon study area differed from all other study areas. There appears to be little or no evidence for temporal variation in adult apparent survival on any of the study areas. Although we did not directly compare fecundity estimates were highly variable among years within all study areas (CV of temporal process variation = 0.672-0.817). Estimates for fecundity among the study populations were Lassen (b = 0.336, SE = 0.083), Eldorado (b = 0.409, SE = 0.087), Sierra (b = 0.284, SE = 0.073), Sequoia and Kings Canyon (b = 0.289, SE = 0.074), and San Bernardino (b = 0.362, SE = 0.038). During most years, the Sierra Nevada populations showed either moderate or poor fecundity. However, 1992 appeared to be an exceptional reproductive year for owls in the Sierra Nevada. In contrast, the San Bernardino population had less variable reproduction (CV of temporal process variation = 0.217), but experienced neither the exceptional reproduction of 1992 nor the extremely poor years that characterized all of the Sierra Nevada study areas. Because fecundity may be influenced by weather patterns, it was possible that the different weather patterns between southern California and the Sierra Nevada accounted for that difference. Except for Eldorado, all estimates for 8t, were <1.0, but none was different from 8 = 1.0 given the 95% confidence i
The relative importance of genetic, demographic, environmental, and catastrophic processes that maintain population bottlenecks has received little consideration. We evaluate the role of these factors in maintaining the Puerto Rican Parrot (Amazona vittata) in a prolonged bottleneck from 1973 through 2000 despite intensive conservation efforts. We first conduct a risk analysis, then examine evidence for the importance of specific processes maintaining the bottleneck using the multiple competing hypotheses approach, and finally integrate these results through a sensitivity analysis of a demographic model using life-stage simulation analysis (LSA) to determine the relative importance of genetic, demographic, environmental, and catastrophic processes on population growth. Annual population growth has been slow and variable (1.0 6 5.2 parrots per year, or an average k?1.05 6 0.19) from 16 parrots (1973) to a high of 40-42 birds (1997-1998). A risk analysis based on population prediction intervals (PPI) indicates great risk and large uncertainty, with a range of 22?83 birds in the 90% PPI only five years into the future. Four primary factors (reduced hatching success due to inbreeding, failure of adults to nest, nest failure due to nongenetic causes, and reduced survival of adults and juveniles) were responsible for maintaining the bottleneck. Egghatchability rates were low (70.6% per egg and 76.8% per pair), and hatchability increased after mate changes, suggesting inbreeding effects. Only an average of 34% of the population nested annually, which was well below the percentage of adults that should have reached an age of first breeding (41-56%). This chronic failure to nest appears to have been caused primarily by environmental and/or behavioral factors, and not by nest-site scarcity or a skewed sex ratio. Nest failure rates from nongenetic causes (i.e., predation, parasitism, and wet cavities) were low (29%) due to active management (protecting nests and fostering captive young into wild nests), diminishing the importance of nest failure as a limiting factor. Annual survival has been periodically reduced by catastrophes (hurricanes), which have greatly constrained population growth, but survival rates were high under non-catastrophic conditions. Although the importance of factors maintaining the Puerto Rican Parrot bottleneck varied throughout the 30-year period of study, we determined their long-term influence using LSA simulations to correlate variation in demographic rates with variation in population growth (k). The bottleneck appears to have been maintained primarily by periodic catastrophes (hurricanes) that reduced adult survival, and secondarily by environmental and/or behavioral factors that resulted in a failure of many adults to nest. The influence of inbreeding through reduced hatching success played a much less significant role, even when additional effects of inbreeding on the production and mortality of young were incorporated into the LSA. Management actions needed to speed recovery include (1) continued nest guarding to minimize the effects of nest failure due to nongenetic causes; (2) creating a second population at another location on the island --a process that was recently initiated--to reduce the chance that hurricane strikes will cause extinction; and (3) determining the causes of the low percentage of breeders in the population and ameliorating them, which would have a large impact on population growth.
This review grew out of our realization that models play an increasingly important role in conservation but are rarely used in the research of most avian biologists. Modelers are creating models that are more complex and mechanistic and that can incorporate more of the knowledge acquired by field biologists. Such models require field biologists to provide more specific information, larger sample sizes, and sometimes new kinds of data, such as habitat-specific demography and dispersal information. Field biologists need to support model development by testing key model assumptions and validating models. The best conservation decisions will occur where cooperative interaction enables field biologists, modelers, statisticians, and managers to contribute effectively. We begin by discussing the general form of ecological models—heuristic or mechanistic, "scientific" or statistical—and then highlight the structure, strengths, weaknesses, and applications of six types of models commonly used in avian conservation: (1) deterministic single-population matrix models, (2) stochastic population viability analysis (PVA) models for single populations, (3) metapopulation models, (4) spatially explicit models, (5) genetic models, and (6) species distribution models. We end by considering their unique attributes, determining whether the assumptions that underlie the structure are valid, and testing the ability of the model to predict the future correctly.
A generalized picture of ice gouge characteristics from shallow inshore depths to the outer shelf at about 60 m of water is presented. Data from recent studies show that the size and quantity of gouging increases in an offshore direction to depths of about 45 m where this trend then reverses and the features decrease in size and quantity as the shelf break is approached. Ice gouges are oriented east-west and this suggests that most gouging is caused by ice approaching from the east, possibly driven by the Beaufort Sea gyre. The most intense gouging occurs in the stamukhi zone, between 20 and 40 m of water, and is caused by a high rate of ice keel production owing to shearing forces between mobile and stable sea ice. Inshore of the stamukhi zone, ice gouging still presents a significant hazard but their greatly decreased size and number make it possible to design against this hazard.
Lesser snow geese ( Chen caerulescens caerulescens ) of the Western Canadian Arctic Population feed intensively for 2-4 weeks on the coastal plain of the Beaufort Sea in Canada and Alaska at the beginning of their autumn migration. Petroleum leasing proposed for the Alaskan portion of the staging area on the Arctic National Wildlife Refuge (ANWR) could affect staging habitats and their use by geese. Therefore we studied availability, distribution, and use by snow geese of tall and russett cotton-grass ( Eriophorum angustifolium and E. russeolum , respectively) feeding habitats on the ANWR. We studied selection of feeding habitats at 3 spatial scales (feeding sites [0.06 m 2 ], feeding patches [ca. 100 m 2 ], and feeding areas [>1 ha]) during 1990-93. We used logistic regression analysis to discriminate differences in soil moisture and vegetation between 1,548 feeding sites where snow geese exploited individual cotton-grass plants and 1,143 unexploited sites at 61 feeding patches in 1990. Feeding likelihood increased with greater soil moisture and decreased where nonforage species were present. We tested the logistic regression model in 1991 by releasing human-imprinted snow geese into 4 10 × 20-m enclosed plots where plant communities had been mapped, habitats sampled, and feeding probabilities calculated. Geese selected more feeding sites per square meter in areas of predicted high quality feeding habitat (feeding probability ≥ 0.6) than in medium (feeding probability = 0.3-0.59) or poor (feeding probability < 0.3) quality habitat (P < 0.0001). Geese increasingly used medium quality areas and spent more time feeding as trials progressed and forage was presumably reduced in high quality habitats. We examined relationships between underground biomass of plants, feeding probability, and surface microrelief at 474 0.06- m 2 sites in 20 thermokarst pits in 1992. Feeding probability was correlated with the percentage of underground biomass composed of cotton-grass (r = 0.56). Feeding probability and relative availability of cotton-grass forage were highest in flooded soils along the ecotone of flooded and upland habitats. In 1992, we also used the logistic regression model to estimate availability of high quality feeding sites on 192 80 × 90-m plots that were randomly located on 24 study areas. A mean of 1.6% of the area sampled in each plot was classified as high quality feeding habitat at 23 of the study areas. Relative availability of high quality sites was highest in troughs, thermokarst pits, and water tracks because saturated soils in those microreliefs were dominated by cotton-grass. Relative availability of high quality sites was lower in saturated soils of basins (low-centered polygons, wet meadows, and strangmoor) because that microrelief was dominated by Carex spp. Most (63%) of the saturated area on the ANWR coastal plain was in basins. We examined distribution of feeding patches relative to microrelief in 49 snow goose feeding areas in 1993. Only 2.5% of the tundra in each feeding area was exploited by snow geese. Snow geese preferentially fed in thermokarst pits, water tracks, and troughs, and avoided basins and uplands. Feeding areas had more thermokarst pit but less basin microrelief than adjacent randomly-selected areas. Thermokarst pits and water tracks occurred most frequently in regions of the coastal plain where geese were observed most often during aerial surveys (1982-93). Microrelief influenced selection of feeding patches and feeding areas and may have affected snow goose distribution on the ANWR. Potential feeding patches were widely distributed but composed a small percentage (≤2.5%) of the tundra landscape and were highly interspersed with less suitable habitat. The Western Canadian Arctic Population probably used a large staging area on the Beaufort Sea coastal plain because snow geese exploited a spatially and temporally heterogeneous resource.
In response to the growing recognition of engineers and decision-makers of the regional effects of earthquake-induced landslides, this paper presents a general approach to conducting seismic landslide zonation, based on the popular Newmark's sliding block analogy for modeling coherent landslides. Four existing models based on the sliding block analogy are compared. The comparison shows that the models forecast notably different levels of slope performance. Considering this discrepancy along with the limitations of static maps as a decision tool, a spatial decision support system (SDSS) for seismic landslide analysis is proposed, which will support investigations over multiple scales for any number of earthquake scenarios and input conditions. Most importantly, the SDSS will allow use of any seismic landslide analysis model and zonation approach. Developments associated with the SDSS will produce an object-oriented model for encapsulating spatial data, an object-oriented specification to allow construction of models using modular objects, and a direct-manipulation, dynamic user-interface that adapts to the particular seismic landslide model configuration.
This paper describes the visualization of spatially distributed water pipeline damage following an earthquake using geographical information systems (GIS). Pipeline damage is expressed as a repair rate (RR). Repair rate contours are developed with GIS by dividing the study area into grid cells (n ?? n), determining the number of particular pipeline repairs in each grid cell, and dividing the number of repairs by the length of that pipeline in each cell area. The resulting contour plot is a two-dimensional visualization of point source damage. High damage zones are defined herein as areas with an RR value greater than the mean RR for the entire study area of interest. A hyperbolic relationship between visual display of high pipeline damage zones and grid size, n, was developed. The relationship is expressed in terms of two dimensionless parameters, threshold area coverage (TAC) and dimensionless grid size (DGS). The relationship is valid over a wide range of different map scales spanning approximately 1,200 km2 for the largest portion of the Los Angeles water distribution system to 1 km2 for the Marina in San Francisco. This relationship can aid GIS users to get sufficiently refined, but easily visualized, maps of damage patterns.
Young Holocene deposits of the Duwamish River valley underlie a highly developed transportation-industrial corridor, extending from the City of Kent to the Elliott Bay-Harbor Island marine terminal facilities. The deposits have been shaped by relative sea-level rise, but also by episodic volcanism and seismicity. A geologic and geotechnical investigation of these river-mouth deposits indicates high initial liquefaction susceptibility during earthquakes, and possibly the potential for unlimited-strain disintegrative flow failure of the delta front.
We studied the dispersal behavior of 1,475 northern spotted owls (Strix occidentalis caurina) during banding and radio-telemetry studies in Oregon and Washington in 1985-1996. The sample included 324 radio-marked juveniles and 1,151 banded individuals (711 juveniles, 440 non-juveniles) that were recaptured or resighted after dispersing from the initial banding location. Juveniles typically left the nest during the last week in May and the first two weeks in June (x?? ?? SE = 8 June ?? 0.53 days, n = 320, range = 15 May-1 July), and spent an average of 103.7 days in the natal territory after leaving the nest (SE = 0.986 days, n = 137, range = 76-147 days). The estimated mean date that juveniles began to disperse was 19 September in Oregon (95% CI = 17-21 September) and 30 September in Washington (95% CI = 25 September-4 October). Mean dispersal dates did not differ between males and females or among years. Siblings dispersed independently. Dispersal was typically initiated with a series of rapid movements away from the natal site during the first few days or weeks of dispersal. Thereafter, most juveniles settled into temporary home ranges in late October or November and remained there for several months. In February-April there was a second pulse of dispersal activity, with many owls moving considerable distances before settling again in their second summer. Subsequent dispersal patterns were highly variable, with some individuals settling permanently in their second summer and others occupying a series of temporary home ranges before eventually settling on territories when they were 2-5 years old. Final dispersal distances ranged from 0.6-111.2 km for banded juveniles and 1.8-103.5 km for radio-marked juveniles. The distribution of dispersal distances was strongly skewed towards shorter distances, with only 8.7% of individuals dispersing more than 50 km. Median natal dispersal distances were 14.6 km for banded males, 13.5 km for radio-marked males, 24.5 km for banded females, and 22.9 km for radio-marked females. On average, banded males and females settled within 4.2 and 7.0 territory widths of their natal sites, respectively. Maximum and final dispersal distances were largely independent of the number of days that juveniles were tracked. Although statistical tests of dispersal direction based on all owls indicated that direction of natal dispersal was non-random, the mean angular deviations and 95% CI's associated with the samples were large, and r-values (vector length) were small. This lead us to conclude that significant test results were the result of large sample size and were not biologically meaningful. Our samples were not large enough to test whether dispersal direction from individual territories was random. In the sample of radio-marked owls, 22% of males and 44% of females were paired at 1 year of age, but only 1.5% of males and 1.6% of females were actually breeding at 1 year of age. At 2 years of age, 68% of males and 77% of females were paired, but only 5.4% of males and 2.6% of females were breeding. In contrast to the radio-marked owls, most juveniles that were banded and relocated at 1 or 2 years of age were paired, although few were breeding. Although recruitment into the territorial population typically occurred when owls were 1-5 years old, 9% of banded juveniles were not recaptured until they were > 5 years old. We suspect that our estimates of age at recruitment of banded owls are biased high because of the likelihood that some individuals were not recaptured in the first year that they entered the territorial population. A minimum of 6% of the banded, non-juvenile owls on our demographic study areas changed territories each year (breeding dispersal). The likelihood of breeding dispersal was higher for females, young owls, owls that did not have a mate in the previous year, and owls that lost their mate from the previous year through death or divorce. Mean and median distances dispersed by adults were
We analyzed and radiocarbon-dated 205 fossil woodrat middens from 14 sites in central and northern Wyoming and adjacent Utah and Montana to document spatiotemporal patterns of Holocene invasion by Utah juniper (Juniperus osteosperma). Holocene migration into central and northern Wyoming and southern Montana from the south proceeded by a series of long-distance dispersal events, which were paced by climate variability and structured by the geographic distribution and connectivity of suitable habitats on the landscape. The migration of Utah juniper into the region involved multiple long-distance dispersal events, ranging from 30 to 135 km. One of the earliest established populations, on East Pryor Mountain in south central Montana, is currently the northernmost population of the species. Establishment by long-distance dispersal of that population and another in the Bighorn Basin occurred during a period of relatively dry climate between 7500 and 5400 years ago. Further expansion of these initial colonizing populations and backfilling to occupy suitable sites to the south was delayed during a wet period from 5400 to 2800 years ago. Development of dry conditions 2800 years ago led to a rapid expansion in which Utah juniper colonized sites throughout its current range. Landscape structure and climate variability play important roles in governing the pattern and pace of natural invasions and deserve close attention in studying and modeling plant invasions, whether exotic or natural.
The West Indian manatee ( Trichechus manatus ) is endangered by human activities throughout its range, including the U.S. Atlantic coast where habitat degradation from coastal development and manatee deaths from watercraft collisions have been particularly severe. We radio-tagged and tracked 78 manatees along the east coast of Florida and Georgia over a 12-year period (1986-1998). Our goals were to characterize the seasonal movements, migratory behavior, and site fidelity of manatees in this region in order to provide information for the development of effective conservation strategies. Most study animals were tracked remotely with the Argos satellite system, which yielded a mean (SD) of 3.7 (1.6) locations per day; all were regularly tracked in the field using conventional radiotelemetry methods. The combined data collection effort yielded >93,000 locations over nearly 32,000 tag-days. The median duration of tracking was 8.3 months per individual, but numerous manatees were tracked over multiple years (max = 6.8 years). Most manatees migrated seasonally over large distances between a northerly warm-season range and a southerly winter range (median one-way distance = 280 km, max = 830 km), but 12% of individuals were resident in a relatively small area (<50 km) year-round. The movements of one adult male spanned >2,300 km of coastline between southeastern Florida and Rhode Island. No study animals journeyed to the Gulf coast of Florida. Regions heavily utilized by tagged manatees included: Fernandina Beach, FL to Brunswick, GA in the warm season; northern Biscayne Bay to Port Everglades, FL in the winter; and central coastal Florida, especially the Banana River and northern Indian River lagoons, in all seasons. Daily travel rate, defined as the distance between successive mean daily locations, averaged 2.5 km (SD = 1.7), but this varied with season, migratory pattern, and sex. Adult males traveled a significantly greater distance per day than did adult females for most of the warm season, which corresponded closely with the principal period of breeding activity, but there was no difference between the sexes in daily travel rate during the winter. The timing of seasonal migrations differed markedly between geographic regions. Most long-distance movements in the southern half of the study area occurred between November and March in response to changing temperatures, whereas most migrations in the northern region took place during the warmer, non-winter months. Manatees left their warm-season range in central Florida in response to cold fronts that dropped water temperatures by an average of 2.0??C over the 24-hr period preceding departure. Water temperature at departure from the warm-season range averaged 19??C, but varied among individuals (16-22??C) and was not related to body size or female reproductive status. The presence of industrial warm-water effluents permitted many manatees to overwinter north of their historic winter range, and for some migrants this delayed autumn migrations and facilitated earlier spring migrations. Southward autumn and northward spring migrations lasted an average of 10 and 15 days at mean rates of 33.5 (SD = 7.6) and 27.3 (SD = 10.5) km/day, respectively. The highest rate of travel during migration was 87 km/day (3.6 km/hr) during winter. Manatees overwintering in southeastern Florida often traveled north during mild weather - sometimes reaching their warm-season range - only to return south again with the next major cold front. Manatees were consistent in their seasonal movement patterns across years and showed strong fidelity, to warm-season and winter ranges. Within a season, individuals usually occupied only 1 or 2 core use areas that encompassed about 90% of daily locations. Most manatees returned faithfully to the same seasonal ranges year after year (median distance between range centers was <5 km between years). Seasonal movements of 4 immature manatees tracked as calves with their mothers
The population status of the Florida black bear ( Ursus americanus floridanus ) is problematic within many portions of its range and its potential listing as a federally threatened species has been the subject of legal debate. We studied Florida black bears in 2 areas in the Okefenokee-Osceola ecosystem in southeast Georgia (i.e.,Okefenokee) and north Florida (i.e., Osceola) from 1995 to 1999 to evaluate relationships between population characteristics, habitat conditions, and human activities. Bears in Okefenokee were hunted and those in Osceola were not. We captured 205 different black bears (124M:81F) 345 times from June 1995 to September 1998. We obtained 13,573 radiolocations from 87 (16M:71F) individual bears during the study. In Okefenokee, black gum ( Nyssa sylvatica ) and saw palmetto ( Serenoa repens ) fruits were the most important foods for bears based on scat analysis. In Osceola, corn from white-tailed deer ( Odocoileus virginianus ) feeders was the most stable food source but saw palmetto was heavily used when available. Corn from deer feeders was not available in Okefenokee. Adult bears in Osceola were 29% heavier than those in Okefenokee ( t 82 = 3.55, P <0.001). The mean annual home-range size for Osceola females ( x̄ =30.3 km 2 ± 4.0 [SE], n =53) varied little seasonally or annually and was almost half that of Okefenokee females (55.9 km 2 ± 6.9, n = 69; Z = –2.47, P = 0.014). In contrast, radiocollared females in Okefenokee expanded their home ranges during years of poor black gum production. That expansion was most apparent between autumn 1998 and 1999, when mean home-range size for Okefenokee females increased from 14.5 km 2 to 78.4 km 2 , respectively, and included a larger proportion of upland areas open to sport hunting. As a result, 5 females were harvested in the Okefenokee study area during the 1999 bear hunting season compared with only 7 harvested from 1996 to 1998. Home ranges of adult female bears were located in areas with disproportionately high loblolly bay ( Gordonia lasianthus ) and gum-bay-cypress ( Taxodium spp.) vegetation associations in Okefenokee and gum-bay-cypress associations in Osceola. The pine vegetation association ranked lower than most other associations within the home ranges of bears in both study areas even though much of the summer and autumn diets of bears included food items found almost exclusively in pine. Sixteen mortalities of radiocollared bears were documented in Okefenokee; hunting accounted for 11 (68.8%) of these deaths. The annual survival rate of radiocollared males in Okefenokee was 0.71 (95% CI = 0.53–0.88) whereas survival of females in Okefenokee was higher ( Z =18.87, P <0.001) at 0.89 (95% CI = 0.83–0.95). The survival rate for females in Osceola was 0.97 (95% CI = 0.92–1.00). Overall, 67 bears (51M:16F) were killed by hunters in the Okefenokee study area from 1995 to 1999. Based on all radiocollared bears in Okefenokee, many of which resided within areas closed to hunting, we estimated an annual harvest rate of 0.22 (95% CI = 0.03–0.37) for males and 0.07 (95% CI = 0.01–0.12) for females. When we excluded those bears that were not in areas open to hunting, however, the annual harvest rate rose to 0.37 (95% CI = 0.07–0.58) for males and 0.39 (95% CI = 0.09–0.58) for females. Following a black gum shortage in autumn 1995, only 1 of 15 radiocollared females in Okefenokee produced cubs in winter 1996. That low reproductive rate was in contrast to winter 1997, which followed heavy black gum production, when 21 of 22 radiocollared females produced cubs. Reproductive output was more consistent in the Osceola study area, with 46 cubs being produced from 8, 5, and 9 litters in 1997, 1998, and 1999, respectively. To estimate population size, we maintained 88 and 94 barbed-wire hair traps during 1999 in the Okefenokee and Osceola study areas, respectively. Using DNA collected at the hair traps, mark–recapture models produced estimates of 71 bears (95% CI = 59–91) in Okefenokee and 44 bears (95% CI = 40–57) in the Osceola study area during 1999. The estimated densities in the Okefenokee and Osceola study areas were 0.12 and 0.14 bears/km 2 , respectively. Alternative density estimates based on the amount of time bears spent within study area boundaries were similar (0.11 and 0.14 bears/km 2 on Okefenokee and Osceola, respectively). We used a population model to estimate the effect of harvest in the Okefenokee bear population. Excluding harvest, bears at Osceola experienced higher average annual population growth (λ = 1.184 ± 0.002) than those at Okefenokee (1.064 ± 0.002; t 18 = 3.93, P = 0.001), most likely due to protection from hunting and higher recruitment. Including the effects of emigration and immigration, we estimated an average annual sustainable harvest at Okefenokee of approximately 9 bears (12.6%), which was slightly less than the average 1995–1999 annual harvest of 9.4. That level of hunting in Okefenokee is sustainable but likely represents the highest exploitation rate in the region. Our mark–recapture data from Osceola suggest a high dispersal rate by subadult bears, and our population modeling data support this hypothesis; we documented bears in Okefenokee that originated from Osceola but not the reverse. We speculate that bears from the interior of the Okefenokee National Wildlife Refuge (ONWR), and to some extent northern Florida, served as a source to the population sink caused by hunting mortality in Okefenokee and in the surrounding Georgia counties. Corn from deer feeders was the most probable reason for smaller home-range sizes and greater body masses and reproductive output at Osceola. Changes in management to eliminate or reduce baiting for deer with corn would negatively affect the Osceola bear population. On Okefenokee, sporadic black gum and palmetto production influenced harvest rates and cub production and, thus, governed bear population dynamics. The U.S. Fish and Wildlife Service (USFWS) concluded in 1998 that listing the Florida black bear as threatened under the Endangered Species Act of 1973 was not warranted. That decision was largely based on the stability and protection afforded to a few subpopulations within the range of the subspecies, which includes the Okefenokee-Osceola subpopulation; our results support that conclusion. However, we suggest that metapopulation processes among the various subpopulations be given greater consideration, with the ultimate goal of managing the sub-species as a unit rather than as an assemblage of independent components. Our study illustrates the importance of travel corridors for maintaining metapopulation processes.
The ramifying effects of top predators on food webs traditionally have been studied within the framework of trophic cascades. Trophic cascades are compelling because they embody powerful indirect effects of predators on primary production. Although less studied, indirect effects of predators may occur via routes that are not exclusively trophic. We quantified how the introduction of foxes onto the Aleutian Islands transformed plant communities by reducing abundant seabird populations, thereby disrupting nutrient subsidies vectored by seabirds from sea to land. We compared soil and plant fertility, plant biomass and community composition, and stable isotopes of nitrogen in soil, plants, and other organisms on nine fox-infested and nine historically fox-free islands across the Aleutians. Additionally, we experimentally augmented nutrients on a fox-infested island to test whether differences in plant productivity and composition between fox-infested and fox-free islands could have arisen from differences in nutrient inputs between island types. Islands with historical fox infestations had soils low in phosphorus and nitrogen and plants low in tissue nitrogen. Soils, plants, slugs, flies, spiders, and bird droppings on these islands had low d15N values indicating that these organisms obtained nitrogen from internally derived sources. In contrast, soils, plants, and higher trophic level organisms on fox-free islands had elevated d15N signatures indicating that they utilized nutrients derived from the marine environment. Furthermore, soil phosphorus (but not nitrogen) and plant tissue nitrogen were higher on fox-free than fox-infested islands. Nutrient subsidized fox-free islands supported lush, high biomass plant communities dominated by graminoids. Fox-infested islands were less graminoid dominated and had higher cover and biomass of low-lying forbs and dwarf shrubs. While d15N profiles of soils and plants and graminoid biomass varied with island size and distance from shore, after accounting for these effects differences between fox-infested and fox-free islands still existed. Fertilization over four years caused a 24-fold increase in graminoid biomass and a shift toward a more graminoid dominated plant community typical of fox-free islands. These results indicate that apex predators can influence plant productivity and composition through complex interaction web pathways involving both top-down forcing and bottom-up nutrient exchanges across systems. ?? 2006 by the Ecological Society of America.