Testing environmental DNA from wolf snow tracks for species, sex, and individual identification – An addendum
No abstract available.
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Geology topics
Research about Superior National Forest
Source-linked reports with geographic coverage including Superior National Forest.
The importance of wilderness to wolf (Canis lupus) survival and cause-specific mortality over 50 years
We assessed the relative importance of wilderness to gray wolf (Canis lupus) population dynamics over 50 years in a population that 1) was long extant (i.e., not reintroduced or recolonized), 2) was not subject to harvest in our study area until recently, and 3) used both wilderness and adjacent, mainly public, non-wilderness. We analyzed the survival of radiocollared wolves (n = 756 collared-wolf tenures) during 1968–2018 in the Superior National Forest, Minnesota, USA, including the Boundary Waters Canoe Area Wilderness. Over 50 years, adult annual survival was 78%. Wolves captured in wilderness tended to exhibit higher survival than those captured in nonwilderness, but the difference was more pronounced during harvest years and post-harvest years when wilderness wolf survival remained relatively high and non-wilderness wolf survival dropped (relative to pre-harvest). During Nov–Apr of pre-harvest years for adults, the natural mortality rate was similar for non-wilderness wolves and wilderness wolves (both 6%), but the anthropogenic mortality rate was higher for non-wilderness wolves than wilderness wolves (7% versus 1%), as was the illegal mortality rate (5% versus 1%). During Nov–Apr of preharvest years, wilderness wolves were less likely to die than non-wilderness wolves (p = 0.042; hazard ratio = 0.59), pups were more likely to die than adults (p = 0.002; hazard ratio = 1.84), and males were less likely to die than females (p = 0.053; hazard ratio = 0.73). Our long-term wolf survival, cause-specific mortality, and hazard results will inform management agencies whenever wolves are delisted, and jurisdiction for them passes to states.
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Use of non-invasive genetics to generate core-area population estimates of a threatened predator in the Superior National Forest, USA
Canada lynx (Lynx canadensis) are found in boreal forests of Canada and Alaska and range southward into the contiguous United States. Much less is understood about lynx in their southern range compared to northern populations. Because lynx are currently listed as threatened under the US Endangered Species Act but have recently been recommended for delisting, information on their southern populations is important for lynx recovery, conservation, and management. We used non-invasive, genetic data collected during lynx snowtracking surveys from 2012-2017 to generate core-area estimates of abundance, trend and density in selected core areas of the Superior National Forest of Minnesota, USA. Lynx abundance estimates averaged 41.8 (SD=14.7, range=24-67) during 2012-2017 in the smaller core areas and averaged 52.3 (SD=8.3, range=43-59) during 2015-2017 in the larger core areas. We found no evidence for a decrease or increase in abundance during either period. Lynx density estimates were approximately 7-10 times lower than densities of lynx in northern populations at the low of the snowshoe hare (Lepus americanus) population cycle. To our knowledge, our results are the first attempt to estimate abundance, trend and density of lynx in Minnesota using non-invasive genetic capture-mark-recapture. Estimates such as ours provide useful benchmarks for future comparisons by providing a context with which to assess 1) potential changes in forest management that may affect lynx recovery and conservation, and 2) possible effects of climate change on the depth, density, and duration of annual snow cover and correspondingly, potential effects on snowshoe hares as well.
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Seasonality of intraspecific mortality by gray wolves
Of 41 adult wolf-killed gray wolves ( Canis lupus ) and 10 probably or possibly killed by wolves from 1968 through 2014 in the Superior National Forest (SNF) in northeastern Minnesota, most were killed in months leading up to and immediately following the breeding season, which was primarily February. This finding is similar to a published sample from Denali National Park, and the seasonality of intraspecific mortality generally parallels the known seasonality of testosterone levels, scent-marking, howling frequency, and general interpack aggression. Males and females were killed in the same proportion as in the population of radiocollared wolves. The annual rate of wolf-killed wolves was not related to the annual wolf density. Our findings tend to support intraspecific mortality of adult wolves as a means to reduce breeding competition and to maintain territories.
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Wolf ( Canis lupus ) generation time and proportion of current breeding females by age
Information is sparse about aspects of female wolf ( Canis lupus ) breeding in the wild, including age of first reproduction, mean age of primiparity, generation time, and proportion of each age that breeds in any given year. We studied these subjects in 86 wolves (113 captures) in the Superior National Forest (SNF), Minnesota (MN), during 1972–2013 where wolves were legally protected for most of the period, and in 159 harvested wolves from throughout MN wolf range during 2012–2014. Breeding status of SNF wolves were assessed via nipple measurements, and wolves from throughout MN wolf range, by placental scars. In the SNF, proportions of currently breeding females (those breeding in the year sampled) ranged from 19% at age 2 to 80% at age 5, and from throughout wolf range, from 33% at age 2 to 100% at age 7. Excluding pups and yearlings, only 33% to 36% of SNF females and 58% of females from throughout MN wolf range bred in any given year. Generation time for SNF wolves was 4.3 years and for MN wolf range, 4.7 years. These findings will be useful in modeling wolf population dynamics and in wolf genetic and dog-domestication studies.
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Predicting tree biomass growth in the temperate-boreal ecotone: is tree size, age, competition or climate response most important?
As global temperatures rise, variation in annual climate is also changing, with unknown consequences for forest biomes. Growing forests have the ability to capture atmospheric CO 2 and thereby slow rising CO 2 concentrations. Forests’ ongoing ability to sequester C depends on how tree communities respond to changes in climate variation. Much of what we know about tree and forest response to climate variation comes from tree-ring records. Yet typical tree-ring datasets and models do not capture the diversity of climate responses that exist within and among trees and species. We address this issue using a model that estimates individual tree response to climate variables while accounting for variation in individuals’ size, age, competitive status, and spatially structured latent covariates. Our model allows for inference about variance within and among species. We quantify how variables influence aboveground biomass growth of individual trees from a representative sample of 15 northern or southern tree species growing in a transition zone between boreal and temperate biomes. Individual trees varied in their growth response to fluctuating mean annual temperature and summer moisture stress. The variation among individuals within a species was wider than mean differences among species. The effects of mean temperature and summer moisture stress interacted, such that warm years produced positive responses to summer moisture availability and cool years produced negative responses. As climate models project significant increases in annual temperatures, growth of species like Acer saccharum , Quercus rubra , and Picea glauca will vary more in response to summer moisture stress than in the past. The magnitude of biomass growth variation in response to annual climate was 92–95% smaller than responses to tree size and age. This means that measuring or predicting the physical structure of current and future forests could tell us more about future C dynamics than growth responses related to climate change alone.
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White-tailed Deer ( Odocoileus virginianus ) fawn risk from Gray Wolf ( Canis lupus ) predation during summer
Little is known about how often various prey animals are at risk of predation by Gray Wolves ( Canis lupus ). We used a system to monitor the presence during the day of two radio-collared Gray Wolves within 2 km of a radio-collared White-tailed Deer ( Odocoileus virginianus ) with a fawn or fawns in August 2013 in the Superior National Forest of northeastern Minnesota. We concluded that the fawn or fawns were at risk of predation by at least one wolf at least daily.
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Evaluation of a formula that categorizes female gray wolf breeding status by nipple size
The proportion by age class of wild Canis lupus (Gray Wolf) females that reproduce in any given year remains unclear; thus, we evaluated the applicability to our long-term (1972–2013) data set of the Mech et al. ( 1993 ) formula that categorizes female Gray Wolf breeding status by nipple size and time of year. We used the formula to classify Gray Wolves from 68 capture events into 4 categories (yearling, adult non-breeder, former breeder, current breeder). To address issues with small sample size and variance, we created an ambiguity index to allow some Gray Wolves to be classed into 2 categories. We classified 20 nipple measurements ambiguously: 16 current or former breeder, 3 former or adult non-breeder, and 1 yearling or adult non-breeder. The formula unambiguously classified 48 (71%) of the nipple measurements; based on supplemental field evidence, at least 5 (10%) of these were incorrect. When used in conjunction with an ambiguity index we developed and with corrections made for classifications involving very large nipples, and supplemented with available field evidence, the Mech et al. ( 1993 ) formula provided reasonably reliable classification of breeding status in wild female Gray Wolves.
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Gray Wolf ( Canis lupus ) dyad monthly association rates by demographic group.
Preliminary data from GPS-collared wolves (Canis lupus) in the Superior National Forest of northeastern Minnesota indicated wolves had low association rates with packmates during summer. However, aerial-telemetry locations of very high frequency (VHF)-radioed wolves in this same area showed high associations among packmates during winter. We analyzed aerial-telemetry-location data from VHF-collared wolves in several packs (n=18 dyads) in this same area from 1994-2012 by month, and found lowest association rates occurred during June. While other studies have found low association among wolf packmates during summer, information on differences in association patterns depending on the wolf associates’ demographics is sparse. During May-July, association rates were greatest for breeding pairs, followed by sibling dyads, and lowest for parent– offspring dyads. Our findings improve our understanding of how individual wolf relationships affect monthly association rates. We highlight some important remaining questions regarding wolf packmate associations.
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How hot is too hot? Live-trapped gray wolf rectal temperatures and 1-year survival
The ability of physically restrained and anesthetized wolves to thermoregulate is lessened and could lead to reduced survival, yet no information is available about this subject. Therefore, we analyzed rectal temperatures related to survival 1 year post-capture from 173 adult (non-pup) gray wolves ( Canis lupus ) captured in modified foot-hold traps for radiocollaring during June–August, 1988–2011, in the Superior National Forest of northeastern Minnesota, USA. The maximum observed rectal temperature (“maxtemp,” ° F, ° C) in each wolf during capture ( x  = 104.0, 40.0; SD = 2.0, 1.1; min. = 95.9, 35.5; max. = 108, 42.2) was not a significant predictor of survival to 1 year post-capture. Although no weather or morphometric variable was a significant predictor of maxtemps, wolves initially anesthetized with ketamine–xylazine rather than telazol®–xylazine averaged higher maxtemps. This information does not fully address possible effects of high body temperatures related to live-capture and handling of wolves, but it does provide a useful waypoint for future assessments of this relationship and a reassurance to wildlife practitioners that the maxtemps observed in our study did not appear to affect 1-year survival.
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Looking for age-related growth decline in natural forests: unexpected biomass patterns from tree rings and simulated mortality
Forest biomass growth is almost universally assumed to peak early in stand development, near canopy closure, after which it will plateau or decline. The chronosequence and plot remeasurement approaches used to establish the decline pattern suffer from limitations and coarse temporal detail. We combined annual tree ring measurements and mortality models to address two questions: first, how do assumptions about tree growth and mortality influence reconstructions of biomass growth? Second, under what circumstances does biomass production follow the model that peaks early, then declines? We integrated three stochastic mortality models with a census tree-ring data set from eight temperate forest types to reconstruct stand-level biomass increments (in Minnesota, USA). We compared growth patterns among mortality models, forest types and stands. Timing of peak biomass growth varied significantly among mortality models, peaking 20–30 years earlier when mortality was random with respect to tree growth and size, than when mortality favored slow-growing individuals. Random or u-shaped mortality (highest in small or large trees) produced peak growth 25–30 % higher than the surviving tree sample alone. Growth trends for even-aged, monospecific Pinus banksiana or Acer saccharum forests were similar to the early peak and decline expectation. However, we observed continually increasing biomass growth in older, low-productivity forests of Quercus rubra, Fraxinus nigra, and Thuja occidentalis. Tree-ring reconstructions estimated annual changes in live biomass growth and identified more diverse development patterns than previous methods. These detailed, long-term patterns of biomass development are crucial for detecting recent growth responses to global change and modeling future forest dynamics.
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Potential increases in natural disturbance rates could offset forest management impacts on ecosystem carbon stocks
Forested ecosystems contain the majority of the world’s terrestrial carbon, and forest management has implications for regional and global carbon cycling. Carbon stored in forests changes with stand age and is affected by natural disturbance and timber harvesting. We examined how harvesting and disturbance interact to influence forest carbon stocks over the Superior National Forest, in northern Minnesota. Forest inventory data from the USDA Forest Service, Forest Inventory and Analysis program were used to characterize current forest age structure and quantify the relationship between age and carbon stocks for eight forest types. Using these findings, we simulated the impact of alternative management scenarios and natural disturbance rates on forest-wide terrestrial carbon stocks over a 100-year horizon. Under low natural mortality, forest-wide total ecosystem carbon stocks increased when 0% or 40% of planned harvests were implemented; however, the majority of forest-wide carbon stocks decreased with greater harvest levels and elevated disturbance rates. Our results suggest that natural disturbance has the potential to exert stronger influence on forest carbon stocks than timber harvesting activities and that maintaining carbon stocks over the long-term may prove difficult if disturbance frequency increases in response to climate change.
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Killing and caching of an adult White-tailed deer, Odocoileus virginianus , by a single Gray Wolf, Canis lupus
A single Gray Wolf ( Canis lupus ) killed an adult male White-tailed Deer ( Odocoileus virginianus ) and cached the intact carcass in 76 cm of snow. The carcass was revisited and entirely consumed between four and seven days later. This is the first recorded observation of a Gray Wolf caching an entire adult deer.
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First record of coccidiosis in Wolves, Canis Lupus
Three 4-month-old Wolf (Canis lupus) pups in the Superior National Forest of Minnesota died during August and September 1997, apparently from coccidiosis. This appears to be the first record of coccidiosis in Wolves.
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Effects of canine parvovirus on gray wolves in Minnesota
Long-term effects of disease on wild animal population demography is not well documented. We studied a gray wolf (Canis lupus) population in a 2,060-km 2 area of Minnesota for 15 years to determine its response to canine parvovirus (CPV). The CPV had little effect ( P > 0.05) on wolf population size while epizootic during 1979-83. However, after CPV became enzootic, percentage of pups captured during summer-fall 1984-93 and changes in subsequent winter wolf numbers were each inversely related to the serological prevalence of CPV in wolves captured during July-November ( r 2 = 0.39 and 0.72, P = 0.05 and < 0.01, respectively). The CPV antibody prevalence in adult wolves increased to 87% in 1993 (r2 = 0.28, P = 0.05). However, because population level remained stable, CPV-induced mortality appeared to compensate for other mortality factors such as starvation. We -predict that the winter wolf population will decline when CPV prevalence in adults consistently exceeds 76%. The CPV may become important in limiting wolf populations.
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Gray wolf density and its association with weights and hematology of pups from 1970 to 1988
We examined weights and hematologic profiles of gray wolf ( Canis lupus ) pups and the associated wolf density in the east-central Superior National Forest of northeastern Minnesota (USA) during 1970 to 1988. We collected weight and hematologic data from 117 pups (57 females, 60 males) during 1 September to 22 November each year. The wolf density (wolves/800 km 2 ) trend was divided into three phases: high (72 ± 7), 1970 to 1975; medium (44 ± 2), 1976 to 1983; and low (27 ± 2), 1984 to 1988. Wolf numbers declined ( P = 0.0001) 39 and 63% from 1970 to 1975 to 1976 to 1983 and from 1970 to 1975 to 1984 to 1988, respectively. Weight was similar between male and female pups and did not vary as wolf density changed. Mean hemoglobin ( P = 0.04), red ( P = 0.0001) and white blood cells ( P = 0.002), mean corpuscular volume, mean corpuscular hemoglobin concentration and mean corpuscular hemoglobin ( P = 0.0001) did differ among the multi-annual phases of changing wolf density. Weight and hematologic data also were compared to values from captive wolf pups. The high, but declining wolf density was associated with macrocytic, normochromic anemia in wolf pups, whereas the lowest density coincided with a hypochromic anemia. Although hematologic values show promise for assessing wolf pup condition and wolf population status, they must be used cautiously until data are available from other populations.
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