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E.D. Forsman

Publications and source records attributed to E.D. Forsman.

8 recordsLinked to original sources

Local weather, regional climate, and annual survival of the northern spotted owl

We used an information-theoretical approach and Cormack—Jolly—Seber models for open populations in program MARK to examine relationships between survival rates of Northern Spotted Owls and a variety of local weather variables and long-term climate variables. In four of the six populations examined, survival was positively associated with wetter than normal conditions during the growing season or high summer temperatures. At the three study areas located at the highest elevations, survival was positively associated with winter temperature but also had a negative or quadratic relation with the number of storms and winter precipitation. A meta-analysis of all six areas combined indicated that annual survival was most strongly associated with phase shifts in the Southern Oscillation and Pacific Decadal Oscillation, which reflect large-scale temperature and precipitation patterns in this region. Climate accounted for a variable amount (1–41%) of the total process variation in annual survival but for more year-to-year variation (3–66%) than did spatial variation among owl territories (0–7%). Negative associations between survival and cold, wet winters and nesting seasons were similar to those found in other studies of the Spotted Owl. The relationships between survival and growing-season precipitation and regional climate patterns, however, had not been reported for this species previously. Climate-change models for the first half of the 21st century predict warmer, wetter winters and hotter, drier summers for the Pacific Northwest. Our results indicate that these conditions could decrease Spotted Owl survival in some areas.

Oregon, Washington

Bald eagles and sea otters in the Aleutian Archipelago: indirect effects of trophic cascades.

Because sea otters ( Enhydra lutris ) exert a wide array of direct and indirect effects on coastal marine ecosystems throughout their geographic range, we investigated the potential influence of sea otters on the ecology of Bald Eagles ( Haliaeetus leucocephalus ) in the Aleutian Islands, Alaska, USA. We studied the diets, productivity, and density of breeding Bald Eagles on four islands during 1993–1994 and 2000–2002, when sea otters were abundant and scarce, respectively. Bald Eagles depend on nearshore marine communities for most of their prey in this ecosystem, so we predicted that the recent decline in otter populations would have an indirect negative effect on diets and demography of Bald Eagles. Contrary to our predictions, we found no effects on density of breeding pairs on four islands from 1993–1994 to 2000–2002. In contrast, diets and diet diversity of Bald Eagles changed considerably between the two time periods, likely reflecting a change in prey availability resulting from the increase and subsequent decline in sea otter populations. The frequency of sea otter pups, rock greenling ( Hexagammus lagocephalus ), and smooth lumpsuckers ( Aptocyclus ventricosus ) in the eagle's diet declined with corresponding increases in Rock Ptarmigan ( Lagopus mutus ), Glaucous-winged Gulls ( Larus glaucescens ), Atka mackerel ( Pleurogrammus monopterygius ), and various species of seabirds during the period of the recent otter population decline. Breeding success and productivity of Bald Eagles also increased during this time period, which may be due to the higher nutritional quality of avian prey consumed in later years. Our results provide further evidence of the wide-ranging indirect effects of sea otter predation on nearshore marine communities and another apex predator, the Bald Eagle. Although the indirect effects of sea otters are widely known, this example is unique because the food-web pathway transcended five species and several trophic levels in linking one apex predator to another.

Ecology

Effects of historical climate change, habitat connectivity, and vicariance on genetic structure and diversity across the range of the Red Tree Vole (Phenacomys longicaudus) in the Pacific Northwest United States

Phylogeographical analyses conducted in the Pacific Northwestern United States have often revealed concordant patterns of genetic diversity among taxa. These studies demonstrate distinct North/South genetic discontinuities that have been attributed to Pleistocene glaciation. We examined phylogeographical patterns of red tree voles (Phenacomys longicaudus) in western Oregon by analysing mitochondrial control region sequences for 169 individuals from 18 areas across the species' range. Cytochrome b sequences were also analysed from a subset of our samples to confirm the presence of major haplotype groups. Phylogenetic network analyses suggested the presence of two haplotype groups corresponding to northern and southern regions of P. longicaudus' range. Spatial genetic analyses (samova and Genetic Landscape Shapes) of control region sequences demonstrated a primary genetic discontinuity separating northern and southern sampling areas, while a secondary discontinuity separated northern sampling areas into eastern and western groups divided by the Willamette Valley. The North/South discontinuity likely corresponds to a region of secondary contact between lineages rather than an overt barrier. Although the Cordilleran ice sheet (maximum a??12 000 years ago) did not move southward to directly affect the region occupied by P. longicaudus, climate change during glaciation fragmented the forest landscape that it inhabits. Signatures of historical fragmentation were reflected by positive associations between latitude and variables such as Tajima's D and patterns associated with location-specific alleles. Genetic distances between southern sampling areas were smaller, suggesting that forest fragmentation was reduced in southern vs. northern regions.

Molecular Ecology

Comment on 'Are survival rates for northern spotted owls biased?'

Loehle et al. recently estimated survival rates from radio-telemetered northern spotted owls (Strix occidentalis caurina (Merriam, 1898)) and suggested that survival rates estimated for this species from capture-recapture studies were negatively biased, which subsequently resulted in the negatively biased estimates of rates of population change (lambda) reported by Anthony et al. (Wildl. Monogr. No. 163, pp. 1-47 (2006)). We argue that their survival estimates were inappropriate for comparison with capture-recapture estimates because (i) the manner in which they censored radio-telemetered individuals had the potential to positively bias their survival estimates, (ii) their estimates of survival were not valid for evaluating bias, and (iii) the size and distribution of their radiotelemetry study areas were sufficiently different from capture-recapture study areas to preclude comparisons. In addition, their inferences of negative bias in rates of population change estimated by Anthony et al. were incorrect and reflected a misunderstanding about those estimators.

Canadian Journal of Zoology

Taxonomic relationships among Phenacomys voles as inferred by cytochrome b

Taxonomic relationships among red tree voles (Phenacomys longicaudus longicaudus, P. l. silvicola), the Sonoma tree vole (P. pomo), the white-footed vole (P. albipes), and the heather vole (P. intermedius) were examined using 664 base pairs of the mitochondrial cytochrome b gene. Results indicate specific differences among red tree voles, Sonoma tree voles, white-footed voles, and heather voles, but no clear difference between the 2 Oregon subspecies of red tree voles (P. l. longicaudus and P. l. silvicola). Our data further indicated a close relationship between tree voles and albipes, validating inclusion of albipes in the subgenus Arborimus. These 3 congeners shared a closer relationship to P. intermedius than to other arvicolids. A moderate association between porno and albipes was indicated by maximum parsimony and neighbor-joining phylogenetic analyses. Molecular clock estimates suggest a Pleistocene radiation of the Arborimus clade, which is concordant with pulses of diversification observed in other murid rodents. The generic rank of Arborimus is subject to interpretation of data.

Journal of Mammalogy

Natal and breeding dispersal of northern spotted owls

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

Wildlife Monographs

Geographic variation and genetic structure in Spotted Owls

We examined genetic variation, population structure, and definition of conservation units in Spotted Owls ( Strix occidentalis ). Spotted Owls are mostly non-migratory, long-lived, socially monogamous birds that have decreased population viability due to their occupation of highly-fragmented late successional forests in western North America. To investigate potential effects of habitat fragmentation on population structure, we used random amplified polymorphic DNA (RAPD) to examine genetic variation hierarchically among local breeding areas , subregional groups , regional groups , and subspecies via sampling of 21 breeding areas (276individuals) among the three subspecies of Spotted Owls. Data from 11 variable bands suggest a significant relationship between geographic distance among local breeding groups and genetic distance (Mantel r = 0.53, P < 0.02) although multi-dimensional scaling of three significant axes did not identify significant grouping at any hierarchical level. Similarly, neighbor-joining clustering of Manhattan distances indicated geographic structure at all levels and identified Mexican Spotted Owls as a distinct clade. RAPD analyses did not clearly differentiate Northern Spotted Owls from California Spotted Owls. Among Northern Spotted Owls, estimates of population differentiation (F ST ) ranged from 0.27 among breeding areas to 0.11 among regions. Concordantly, within-group agreement values estimated via multi-response permutation procedures of Jaccard's distances ranged from0.22 among local sites to 0.11 among regions. Pairwise comparisons of F ST and geographic distance within regions suggested only the Klamath region was in equilibrium with respect to gene flow and genetic drift. Merging nuclear data with recent mitochondrial data provides support for designation of an Evolutionary Significant Unit for Mexican Spotted Owls and two overlapping Management Units for Northern and California Spotted Owls.

Conservation Genetics

Spatial and temporal variation in diets of Spotted Owls in Washington

We studied diets of Northern Spotted Owls (Strix occidentalis caurina) in three different regions of Washington State during 1983-96. Northern flying squirrels (Glaucomys sabrinus) were the most important prey in most areas, comprising 29-54% of prey numbers and 45-59% of prey biomass. Other important prey included snowshoe hares (Lepus americanus), bushy-tailed woodrats (Neoloma cinerea), boreal red-backed voles (Clethrionomys gapperi), and mice (Peromyscus maniculatus, P. oreas). Non-mammalian prey generally comprised less than 15% of prey numbers and biomass. Mean prey mass was 111.4 ?? 1.5 g on the Olympic Peninsula, 74.8 ?? 2.9 g in the Western Cascades, and 91.3 ?? 1.7 g in the Eastern Cascades. Diets varied among territories, years, and seasons. Annual variation in diet was characterized by small changes in relative occurrence of different prey types rather than a complete restructuring of the diet. Predation on snowshoe hares was primarily restricted to small juveniles captured during spring and summer. Mean prey mass did not differ between nesting and nonnesting owls in 19 of 21 territories examined. However, the direction of the difference was positive in 15 of the 21 cases (larger mean for nesting owls), suggesting a trend toward larger prey in samples collected from nesting owls. We suggest that differences in diet among years, seasons, and territories are probably due primarily to differences in prey abundance. However, there are other factors that could cause such differences, including individual variation in prey selection, variation in the timing of pellet collections, and variation in prey accessibility in different cover types. ?? 2001 The Raptor Research Foundation, Inc.

Journal of Raptor Research