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Although sampling the five tallest young aspen in a stand is useful for detecting the occurrence of any aspen recruitment, this technique overestimates the population response of aspen to wolf reintroduction. Our original conclusion that random sampling described a trophic cascade that was weaker than the one described by non‐random sampling is unchanged. [ABSTRACT FROM AUTHOR]

Genetic diversity is theorized to decrease in populations closer to a species' range edge, where habitat may be suboptimal. Generalist species capable of long‐range dispersal may maintain sufficient gene flow to counteract this, though the presence of significant barriers to dispersal (e.g., large water bodies, human‐dominated landscapes) may still lead to, and exacerbate, the edge effect. We used microsatellite data for 2421 gray wolves (Canis lupus) from 24 subpopulations (groups) to model how allelic richness and expected heterozygosity varied with mainland–island position and two measures of range edge (latitude and distance from range center) across >7.3 million km2 of northern North America. We expected low genetic diversity both at high latitudes, due to harsh environmental conditions, and on islands, but no change in diversity with distance to the range center due to the species' exceptional dispersal ability and favorable conditions in far eastern and western habitats. We found that allelic richness and expected heterozygosity of island groups were measurably less than that of mainland groups, and that these differences increased with the island's distance to the species' range center in the study area. Our results demonstrate how multiple axes of geographic isolation (distance from range center and island habitation) can act synergistically to erode the genetic diversity of wide‐ranging terrestrial vertebrate populations despite the counteracting influence of long‐range dispersal ability. These findings emphasize how geographic isolation is a potential threat to the genetic diversity and viability of terrestrial vertebrate populations even among species capable of long‐range dispersal. [ABSTRACT FROM AUTHOR]

Although it is well established that density dependence drives changes in organismal abundance over time, relatively little is known about how density dependence affects variation in abundance over space. We tested the hypothesis that spatial trade‐offs between food and safety can change the drivers of population distribution, caused by opposing patterns of density‐dependent habitat selection (DDHS) that are predicted by the multidimensional ideal free distribution. We addressed this using winter aerial survey data of northern Yellowstone elk (Cervus canadensis) spanning four decades. Supporting our hypothesis, we found positive DDHS for food (herbaceous biomass) and negative DDHS for safety (openness and roughness), such that the primary driver of habitat selection switched from food to safety as elk density decreased from 9.3 to 2.0 elk/km2. Our results demonstrate how population density can drive landscape‐level shifts in population distribution, confounding habitat selection inference and prediction and potentially affecting community‐level interactions. [ABSTRACT FROM AUTHOR]

Understanding trophic cascades in terrestrial wildlife communities is a major challenge because these systems are difficult to sample properly. We show how a tradition of non‐random sampling has confounded this understanding in a textbook system (Yellowstone National Park) where carnivore [Canis lupus (wolf)] recovery is associated with a trophic cascade involving changes in herbivore [Cervus canadensis (elk)] behaviour and density that promote plant regeneration. Long‐term data indicate a practice of sampling only the tallest young plants overestimated regeneration of overstory aspen (Populus tremuloides) by a factor of 4–7 compared to random sampling because it favoured plants taller than the preferred browsing height of elk and overlooked non‐regenerating aspen stands. Random sampling described a trophic cascade, but it was weaker than the one that non‐random sampling described. Our findings highlight the critical importance of basic sampling principles (e.g. randomisation) for achieving an accurate understanding of trophic cascades in terrestrial wildlife systems. [ABSTRACT FROM AUTHOR]

Temporal fluctuations in growth rates can arise from both variation in age‐specific vital rates and temporal fluctuations in age structure (i.e. the relative abundance of individuals in each age‐class). However, empirical assessments of temporal fluctuations in age structure and their effects on population growth rate are relatively rare. Most research has focused on understanding the contribution of changing vital rates to population growth rates and these analyses routinely assume that: (a) populations have stable age distributions, (b) environmental influences on vital rates and age structure are stationary (i.e. the mean and/or variance of these processes does not change over time), and (c) dynamics are independent of density.Here we quantified fluctuations in age structure and assessed whether they were stationary for four populations of free‐ranging vertebrates: moose (observed for 48 years), elk (15 years), tawny owls (15 years) and grey wolves (17 years). We also assessed the extent that fluctuations in age structure were useful for predicting annual population growth rates using models which account for density dependence.Fluctuations in age structure were of a similar magnitude to fluctuations in abundance. For three populations (moose, elk, owls), the mean and the skew of the age distribution fluctuated without stabilizing over the observed time periods. More precisely, the sample variance (interannual variance) of age structure indices increased with the length of the study period, which suggests that fluctuations in age structure were non‐stationary for these populations – at least over the 15‐ to 48‐year periods analysed.Fluctuations in age structure were associated with population growth rate for two populations. In particular, population growth varied from positive to negative for moose and from near zero to negative for elk as the average age of adults increased over its observed range.Non‐stationarity in age structure may represent an important mechanism by which abundance becomes non‐stationary – and therefore difficult to forecast – over time‐scales of concern to wildlife managers. Overall, our results emphasize the need for vertebrate populations to be modelled using approaches that consider transient dynamics and density dependence and that do not rely on the assumption that environmental processes are stationary. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

The extent to which prey space use actively minimizes predation risk continues to ignite controversy. Methodological reasons that have hindered consensus include inconsistent measurements of predation risk, biased spatiotemporal scales at which responses are measured and lack of robust null expectations.We addressed all three challenges in a comprehensive analysis of the spatiotemporal responses of adult female elk (Cervus elaphus) to the risk of predation by wolves (Canis lupus) during winter in northern Yellowstone, USA.We quantified spatial overlap between the winter home ranges of GPS‐collared elk and three measures of predation risk: the intensity of wolf space use, the distribution of wolf‐killed elk and vegetation openness. We also assessed whether elk varied their use of areas characterized by more or less predation risk across hours of the day, and estimated encounter rates between simultaneous elk and wolf pack trajectories. We determined whether observed values were significantly lower than expected if elk movements were random with reference to predation risk using a null model approach.Although a small proportion of elk did show a tendency to minimize use of open vegetation at specific times of the day, overall we highlight a notable absence of spatiotemporal response by female elk to the risk of predation posed by wolves in northern Yellowstone.Our results suggest that predator–prey interactions may not always result in strong spatiotemporal patterns of avoidance. [ABSTRACT FROM AUTHOR]