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Geology topics

E.D. Forsman

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

13 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↗

Population trends in northern spotted owls: Associations with climate in the Pacific Northwest

We used reverse time capture-mark-recapture models to describe associations between rate of population change (??) and climate for northern spotted owls (Strix occidentalis caurina) at six long-term study areas in Washington and Oregon, USA. Populations in three of six areas showed strong evidence of declining populations, while populations in two additional areas were likely declining as well. At four areas, ?? was positively associated with wetter-than-normal conditions during the growing season, which likely affects prey availability. Lambda was also negatively associated with cold, wet winters and nesting seasons, and the number of hot summer days. The amount of annual variation in ?? accounted for by climate varied across study areas (3-85%). Rate of population change was more sensitive to adult survival than to recruitment; however, there was considerable variation among years and across study areas for all demographic rates. While annual survival was more closely related to regional climate conditions, recruitment was often associated with local weather. In addition to climate, declines in recruitment at four of six areas were associated with increased presence of barred owls. Climate change models predict warmer, wetter winters and hotter, drier summers for the Pacific Northwest in the first half of the 21st century. Our results indicate that these conditions have the potential to negatively affect annual survival, recruitment, and consequently population growth rates for northern spotted owls. ?? 2010 Elsevier Ltd.

Biological Conservation↗

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↗

Status and trends in demography of northern spotted owls, 1985-2003

We analyzed demographic data from northern spotted owls (Strix occidentalis caurina) from 14 study areas in Washington, Oregon, and California for 1985-2003. The purpose of our analyses was to provide an assessment of the status and trends of northern spotted owl populations throughout most of their geographic range. The 14 study areas made up approximately 12% of the range of the subspecies and included federal, tribal, private, and mixed federal and private lands. The study areas also included all the major forest types that the subspecies inhabits. The analyses followed rigorous protocols that were developed a priori and were the result of extensive discussions and consensus among the authors. Our primary objectives were to estimate fecundity, apparent survival (??), and annual rate of population change (??) and to determine if there were any temporal trends in these population parameters. In addition to analyses of data from individual study areas, we conducted 2 meta-analyses on each demographic parameter. One meta-analysis was conducted on all 14 areas, and the other was restricted to the 8 areas that constituted the Effectiveness Monitoring Plan for northern spotted owls under the Northwest Forest Plan. The average number of years of reproductive data per study area was 14 (range = 5-19), and the average number of recapture occasions per study area was 13 (range = 4-18). Only 1 study area had <12 years of data. Our results were based on 32,054 captures and resightings of 11,432 banded individuals for estimation of survival and 10,902 instances in which we documented the number of young produced by territorial females. The number of young fledged (NYF) per territorial female was analyzed by testing a suite of a priori models that included (1) effects of age, (2) linear or quadratic time trends, (3) presence of barred owls (Strix varia) in spotted owl territories, and (4) an even-odd year effect. The NYF varied among years on most study areas with a biennial cycle of high reproduction in even-numbered years and low reproduction in odd-numbered years. These cyclic fluctuations did not occur on all study areas, and the even-odd year effect waned during the last 5 years of the study. Fecundity was highest for adults (x??=0.372, SE=0.029), lower for 2-year-olds (x??=0.208, SE=0.032), and very low for 1-year-olds (x??=0.074, SE = 0.029). Fecundity was stable over time for 6 areas (Rainier, Olympic, Warm Springs, H. J. Andrews, Klamath, and Marin), declining for 6 areas (Wenatchee, Cle Elum, Oregon Coast Range, Southern Oregon Cascades, Northwest California, and Simpson), and slightly increasing for 2 areas (Tyee, Hoopa). We found little association between NYF and the proportion of northern spotted owl territories where barred owls were detected, although results were suggestive of a negative effect of barred owls on the Wenatchee and Olympic study areas. The meta-analysis on fecundity indicated substantial annual variability with no increasing or decreasing trends. Fecundity was highest in the mixed-conifer region of eastern Washington (x??=0.560, SE=0.041) and lowest in the Douglas-fir (Pseudotsuga menziesii) region of the Oregon coast (x??=0.306, SE=0.039). We used Cormack-Jolly-Seber open population models and Program MARK to estimate apparent survival rates of owls >1 year old. We found no differences in apparent survival rates between sexes except for 1 area (Marin), which had only 6 years of data. Estimates of apparent survival from individual study areas indicated that there were differences among age classes with adults generally having higher survival than 1- and 2-year-olds. Apparent survival rates ranged from 0.750 (SE=0.026) to 0.886 (SE=0.010) for adults, 0.626 (SE=0.073) to 0.886 (SE=0.010) for 2-year-olds, and 0.415 (SE=0.111) to 0.860 (SE=0.017) for 1-year-olds. These estimates were comparable to survival rates from previous studies on the subspecies. We found evidence for negative time trends in survival rate

Wildlife Monographs↗

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↗

Genetic identification of spotted owls, barred owls, and their hybrids: Legal implications of hybrid identity

Recent population expansion of Barred Owls ( Strix varia) into western North America has led to concern that they may compete with and further harm the Northern Spotted Owl ( S. occidentalis caurina), which is already listed as threatened under the U.S. Endangered Species Act (ESA). Because they hybridize, there is a legal need under the ESA for forensic identification of both species and their hybrids. We used mitochondrial control-region DNA and amplified fragment-length polymorphism (AFLP) analyses to assess maternal and biparental gene flow in this hybridization process. Mitochondrial DNA sequences (524 base pairs) indicated large divergence between Barred and Spotted Owls (13.9%). Further, the species formed two distinct clades with no signs of previous introgression. Fourteen diagnostic AFLP bands also indicated extensive divergence between the species, including markers differentiating them. Principal coordinate analyses and assignment tests clearly supported this differentiation. We found that hybrids had unique genetic combinations, including AFLP markers from both parental species, and identified known hybrids as well as potential hybrids with unclear taxonomic status. Our analyses corroborated the findings of extensive field studies that most hybrids genetically sampled resulted from crosses between female Barred Owls and male Spotted Owls. These genetic markers make it possible to clearly identify these species as well as hybrids and can now be used for research, conservation, and law enforcement. Several legal avenues may facilitate future conservation of Spotted Owls and other ESA-listed species that hybridize, including the ESA similarity-of-appearance clause (section 4[e]) and the Migratory Bird Treaty Act. The Migratory Bird Treaty Act appears to be the most useful route at this time.

Conservation Biology↗

Subspecific relationships and genetic structure in the spotted owl

Hierarchical genetic structure was examined in the three geographically-defined subspecies of spotted owl (Strix occidentalis) to define relationships among subspecies and quantify variation within and among regional and local populations. Sequences (522 bp) from domains I and II of the mitochondrial control region were analyzed for 213 individuals from 30 local breeding areas. Results confirmed significant differences between northern spotted owls and the other traditional geographically defined subspecies but did not provide support for subspecific level differences between California and Mexican spotted owls. Divergence times among subspecies estimated with a 936 bp portion of the cytochrome b gene dated Northern and California/Mexican spotted owl divergence time to 115,000–125,000 years ago, whereas California/Mexican spotted owl divergence was estimated at 15,000 years ago. Nested clade analyses indicated an association between California spotted owl and Mexican spotted owl haplotypes, implying historical contact between the two groups. Results also identified a number of individuals geographically classified as northern spotted owls ( S. o. caurina ) that contained haplotypes identified as California spotted owls ( S. o. caurina ). Among all northern spotted owls sampled (n=131), 12.9% contained California spotted owl haplotypes. In the Klamath region, which is the contact zone between the two subspecies, 20.3% (n=59) of owls were classified as California spotted owls. The Klamath region is a zone of hybridization and speciation for many other taxa as well. Analyses of population structure indicated gene flow among regions within geographically defined subspecies although there was significant differentiation among northern and southern regions of Mexican spotted owls. Among all areas examined, genetic diversity was not significantly reduced except in California spotted owls where the southern region consists of one haplotype. Our results indicate a stable contact zone between northern and California spotted owls, maintaining distinct subspecific haplotypes within their traditional ranges. This supports recovery efforts based on the traditional subspecies designation for the northern spotted owl. Further, although little variation was found between California and Mexican spotted owls, we suggest they should be managed separately because of current isolation between groups.

Conservation Genetics↗

Diets and foraging behavior of northern Spotted Owls in Oregon

We describe local, regional, and annual variation in diets of northern Spotted Owls (Strix occidentalis caurina) in Oregon based on 24 497 prey collected at 1118 owl territories in 1970-2003. The sample included 91.5% mammals, 4.3% birds, 4.1% insects, and 0.1% other prey. The diet included ???131 species, including 49 mammals, 41 birds, 3 reptiles, 1 frog, 1 crayfish, 1 scorpion, 2 snails, and 33 species of insects. On average, 91.9 ?? 0.3% (SE) of prey in the diet were nocturnal animals, 3.3 ?? 0.2% were diurnal, and 4.8 ?? 0.2% were active both day and night. Of the prey captured, 50.5 ?? 0.8% were arboreal, 18.7 ?? 0.7% were scansorial, 4.8 ?? 0.2% were aerial, and 26.0 = 0.7% were terrestrial. Mean mass of prey was 116.6 ?? 6.5 g. Diets varied among owl territories, geographic regions, and years; but were generally dominated by four to six species of nocturnal mammals, including northern flying squirrels (Glaucomys sabrinus), woodrats (Neotoma fuscipes and N. cinerea), red tree voles (Arborimus longicaudus), western red-backed voles (Clethrionomys californicus), deer mice (Peromyscus maniculatus), or gophers (Thomomys spp.). Estimates of dietary evenness were low, indicating diets dominated by a few species of mammals. Forest management practices that produce healthy populations of arboreal and scansorial mammals such as flying squirrels, woodrats, and red tree voles should benefit northern Spotted Owls in Oregon and Washington. ?? 2004 The Raptor Research Foundation, Inc.

Journal of Raptor Research↗

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↗