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Status review of the Marbled Murrelet (Brachyramphus marmoratus) in Alaska and British Columbia

The Marbled Murrelet ( Brachyramphus marmoratus ) is a small, diving seabird inhabiting inshore waters of the Northeastern Pacific Ocean. This species feeds on small, schooling fishes and zooplankton, and nests primarily on the moss-covered branches of large, old-growth conifers, and also, in some parts of its range, on the ground. We reviewed existing information on this species to evaluate its current status in the northern part of its range—Alaska (U.S.) and British Columbia (Canada). Within the southern part of its range (Washington, Oregon, and California, U.S.), the Marbled Murrelet was listed as a threatened species under the Endangered Species Act (ESA) in 1993, and the U.S. Fish and Wildlife Service (USFWS) needed information on the species throughout its range for ESA deliberations. We compiled published information on the conservation status, population biology, foraging ecology, population genetics, population status and trends, demography, marine and nesting habitat characteristics, threats, and ongoing conservation efforts for Marbled Murrelets in Alaska and British Columbia. We conducted a new genetic study using samples from a segment of the range that had not been included in previous studies (Washington, Oregon) and additional nuclear intron and microsatellite markers. We also analyzed available at-sea survey data from several locations for trend. To understand the reasonableness of the empirical trend data, we developed demographic models incorporating stochasticity to discern what population trends were possible by chance. The genetic studies substantially confirmed previous findings on population structure in the Marbled Murrelet. Our present work finds three populations: (1) one comprising birds in the central and western Aleutian Islands; (2) one comprising birds in central California; and (3) one comprising birds within the center of the range from the eastern Aleutians to northern California. Our knowledge of genetic structure within this central population is limited and it requires additional study. Compiling available abundance information, we estimated that in the recent past, Marbled Murrelets in Alaska numbered on the order of 1 million birds. We were unable to generate a similar estimate for historical population size in British Columbia. Using trend information from at-sea surveys spanning a wide geographic range in Alaska, murrelet numbers declined significantly at five of eight trend sites at annual rates of -5.4 to -12.7 percent since the early 1990s. Applying these rates of decline to the historical population estimate, the current murrelet population in Alaska is projected to be on the order of 270,000 birds. This represents an overall population decline of about 70 percent during the past 25 years. In British Columbia, available trend data indicate that murrelet populations there have experienced similar declines. We updated a recent (2002) population estimate for British Columbia, concluding that there are now between 54,000 and 92,000 murrelets in British Columbia. The rates of decline we observed are within, but at the high end of, a range of rates expected by chance. Given that declines were estimated for sites over essentially the entire northern range of the species, there is cause for concern about the species’ status. In their marine habitats, Marbled Murrelets overlap with salmon (Oncorhynchus sp.) gillnetting operations in British Columbia and in Alaska (especially in Prince William Sound and Southeast Alaska), and annual bycatch mortality is likely in the low thousands per year, although bycatch rates are difficult to measure. The species’ inshore distribution coincides with high levels of vessel traffic and makes them especially vulnerable to both chronic oil pollution and to catastrophic spills (e.g., the 1989 Exxon Valdez oil spill [EVOS] in south-central Alaska, which is estimated to have killed 12,000 to 15,000 murrelets). In their forested nesting habitats, Marbled Murrelets have lost about 15 percent of their suitable nesting habitat in Southeast Alaska, and 33 to 49 percent in British Columbia, from industrial-scale logging within the past half century. Increased predation also may be a threat to murrelet populations, related to fragmentation and edge effects from logging and development, and recent population increases observed for some important murrelet predators, including Bald Eagles ( Haliaeetus leucocephalus ), Common Ravens ( Corvus corax ), and Steller’s Jays ( Cyanocitta stelleri ). Nesting habitat losses cannot explain the declines observed in areas where industrial logging has not occurred on a large scale (e.g., Prince William Sound) or at all (Glacier Bay). The apparent change in population size and rates of decline reported for the Marbled Murrelet are large, and we therefore considered alternative explanations and precedents for changes of similar magnitude in other marine wildlife populations in the Northeastern Pacific Ocean. The declines are likely real, and related to combined and cumulative effects from climate-related changes in the marine ecosystem (most likely the 1977 regime shift) and human activities (logging, gillnet bycatch, oil pollution). Much uncertainty about the decline could be alleviated by continuing to repeat boat surveys in Prince William Sound and lower Cook Inlet, and by repeating the boat survey of Southeast Alaska that was conducted in 1994. This survey used a statistically sound design and covered the region that has been and likely remains the center of the species’ abundance. Important questions remain to be addressed about methods for measuring population status and change, adult mortality (major sources, density dependence, seasonal concordance), and the movements of wintering populations.

Alaska, British Columbia↗

Ecology of Florida black bears in the Okefenokee-Osceola ecosystem

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 &plusmn; 4.0 [SE], n =53) varied little seasonally or annually and was almost half that of Okefenokee females (55.9 km 2 &plusmn; 6.9, n = 69; Z = &ndash;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&ndash;0.88) whereas survival of females in Okefenokee was higher ( Z =18.87, P <0.001) at 0.89 (95% CI = 0.83&ndash;0.95). The survival rate for females in Osceola was 0.97 (95% CI = 0.92&ndash;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&ndash;0.37) for males and 0.07 (95% CI = 0.01&ndash;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&ndash;0.58) for males and 0.39 (95% CI = 0.09&ndash;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&ndash;recapture models produced estimates of 71 bears (95% CI = 59&ndash;91) in Okefenokee and 44 bears (95% CI = 40&ndash;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 (&lambda; = 1.184 &plusmn; 0.002) than those at Okefenokee (1.064 &plusmn; 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&ndash;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&ndash;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.

Florida, Georgia↗