Variable strategies to solve risk-reward tradeoffs in carnivore communities.

Mesopredator release theory suggests that dominant predators suppress subordinate carnivores and ultimately shape community dynamics, but the assumption that subordinate species are only negatively affected ignores the possibility of facilitation through scavenging. We examined the interplay within a carnivore community consisting of cougars, coyotes, black bears, and bobcats using contemporaneous Global Positioning System telemetry data from 51 individuals; diet analysis from 972 DNA-metabarcoded scats; and data from 128 physical investigations of cougar kill sites, 28 of which were monitored with remote cameras. Resource provisioning from competitively dominant cougars to coyotes through scavenging was so prolific as to be an overwhelming determinant of coyote behavior, space use, and resource acquisition. This was evident via the strong attraction of coyotes to cougar kill sites, frequent scavenging of cougar-killed prey, and coyote diets that nearly matched cougars in the magnitude of ungulate consumption. Yet coyotes were often killed by cougars and used space to minimize encounters, complicating the fitness benefits gained from scavenging. We estimated that 23% (95% CI: 8 to 55%) of the coyote population in our study area was killed by cougars annually, suggesting that coyote interactions with cougars are a complex behavioral game of risk and reward. In contrast, we found no indication that bobcat space use or diet was influenced by cougars. Black bears avoided cougars, but there was no evidence of attraction to cougar kill sites and much lower levels of ungulate consumption and carcass visitation than for coyotes. Interspecific interactions among carnivores are multifaceted, encompassing both suppression and facilitation.

Density-dependent changes in elk resource selection over successional time scales following forest disturbance.

There is an increasing need to understand how animals respond to modifications of their habitat following landscape-scale disturbances such as wildfire or timber harvest. Such disturbances can promote increased use by herbivores due to changes in plant community structure that improve forage conditions, but can also cause avoidance if other habitat functions provided by cover are substantially reduced or eliminated. Quantifying the total effects of these disturbances, however, is challenging because they may not fully be apparent unless observed at successional timescales. Further, the effects of disturbances that improve habitat quality may be density dependent, such that the benefits are (1) less valuable to high-density populations because the per-capita benefits are reduced when shared among more users or, alternatively, (2) more valuable to animals living in high densities because resources may be more depleted from the greater intraspecific competition. We used 30 years of telemetry data on elk occurring at two distinct population densities to quantify changes in space use at diel, monthly, and successional timescales following timber harvest. Elk selected logged areas at night only, with selection strongest during midsummer, and peak selection occurring 14 years post harvest, but persisting for 26-33 years. This pattern of increased selection at night following a reduction in overhead canopy cover is consistent with elk exploiting improved nutritional conditions for foraging. The magnitude of selection for logged areas was 73% higher for elk at low population density, consistent with predictions from the ideal free distribution. Yet elk avoided these same areas during daytime for up to 28 years post logging and instead selected untreated forest, suggesting a role for cover to meet other life history requirements. Our results demonstrate that while landscape-scale disturbances can lead to increased selection by large herbivores and suggest that the improvement in foraging conditions can persist over short-term successional timescales, the magnitude of the benefits may not be equal across population densities. Further, the enduring avoidance of logging treatments during the daytime indicates a need for structurally intact forests and suggests that a mosaic of forest patches of varying successional stages and structural completeness is likely to be the most beneficial to large herbivores.

Evaluating and integrating spatial capture-recapture models with data of variable individual identifiability.

Spatial capture-recapture (SCR) models have become the preferred tool for estimating densities of carnivores. Within this family of models are variants requiring identification of all individuals in each encounter (SCR), a subset of individuals only (generalized spatial mark-resight, gSMR), or no individual identification (spatial count or spatial presence-absence). Although each technique has been shown through simulation to yield unbiased results, the consistency and relative precision of estimates across methods in real-world settings are seldom considered. We tested a suite of models ranging from those only requiring detections of unmarked individuals to others that integrate remote camera, physical capture, genetic, and global positioning system (GPS) data into a hybrid model, to estimate population densities of black bears, bobcats, cougars, and coyotes. For each species, we genotyped fecal DNA collected with detection dogs during a 20-d period. A subset of individuals from each species was affixed with GPS collars bearing unique markings and resighted by remote cameras over 140 d contemporaneous with scat collection. Camera-based gSMR models produced density estimates that differed by <10% from genetic SCR for bears, cougars, and coyotes once important sources of variation (sex or behavioral status) were controlled for. For bobcats, SCR estimates were 33% higher than gSMR. The cause of the discrepancies in estimates was likely attributable to challenges designing a study compatible for species with disparate home range sizes and the difficulty of collecting sufficient data in a timeframe in which demographic closure could be assumed. Unmarked models estimated densities that varied greatly from SCR, but estimates became more consistent in models wherein more individuals were identifiable. Hybrid models containing all data sources exhibited the most precise estimates for all species. For studies in which only sparse data can be obtained and the strictest model assumptions are unlikely to be met, we suggest researchers use caution making inference from models lacking individual identity. For best results, we further recommend the use of methods requiring at least a subset of the population is marked and that multiple data sets are incorporated when possible.

Fire history influences large-herbivore behavior at circadian, seasonal, and successional scales.

Recurrent environmental changes often prompt animals to alter their behavior leading to predictable patterns across a range of temporal scales. The nested nature of circadian and seasonal behavior complicates tests for effects of rarer disturbance events like fire. Fire can dramatically alter plant community structure, with important knock-on effects at higher trophic levels, but the strength and timing of fire's effects on herbivores remain unclear. We combined prescribed fire treatments with fine-scale location data to quantify herbivore responses to fire across three temporal scales. Between 2001 and 2003, 26 stands of fir (Abies spp.) and Douglas-fir (Pseudotsuga menziesii) were thinned and burned; 27 similar stands were left untreated as experimental controls. Analyzing female elk (Cervus canadensis) locations across 21 yr (1996-2016), we found crepuscular, seasonal, and successional shifts in behavioral responses to fire. Elk displayed "commuting" behavior, avoiding burns during the day, but selecting them at night. Elk selection for burns was strongest in early summer and the relative probability of elk using burns peaked quickly (5 yr post burn) before gradually returning to pre-treatment levels (15 yr post burn). Our results demonstrate that fire history has complex, persistent effects on herbivore behavior, and suggest that herbivores benefit from heterogeneous landscapes containing a range of successional stages.

A seasonal pulse of ungulate neonates influences space use by carnivores in a multi-predator, multi-prey system.

The behavioral mechanisms by which predators encounter prey are poorly resolved. In particular, the extent to which predators engage in active search for prey versus incidentally encountering them has not been well studied in many systems and particularly not for neonate prey during the birth pulse. Parturition of many large herbivores occurs during a short and predictable temporal window in which young are highly vulnerable to predation. Our study aims to determine how a suite of carnivores responds to the seasonal pulse of newborn ungulates using contemporaneous global positioning system (GPS) locations of four species of predators and two species of prey. We used step-selection functions to assess whether coyotes, cougars, black bears, and bobcats encountered parturient adult female ungulates more often than expected by chance in a low-density population of mule deer and a high-density population of elk. We then assessed whether the carnivore species that encountered parturient prey more often than expected by chance did so by shifting their habitat use toward areas with a high probability of encountering neonates. None of the four carnivore species encountered GPS-collared parturient mule deer more often than expected by chance. By contrast, we determined that cougar and male bear movements positioned them in the proximity of GPS-collared parturient elk more often than expected by chance which may provide evidence of searching behavior. Although both male bears and cougars exhibited behavior consistent with active search for neonates, only male bears used elk parturition habitat in a way that dynamically tracked the phenology of the elk birth pulse suggesting that maximizing encounters with juvenile elk was a motivation when selecting resources. Our results suggest that there is high interspecific and intersexual variability in foraging strategies among large mammalian predators and their prey.

Acidimicrobium ferrooxidans gen. nov., sp. nov.: mixed-culture ferrous iron oxidation with Sulfobacillus species.

A new species of ferrous-iron-oxidizing, moderately thermophilic, acidophilic bacteria, Acidimicrobium ferrooxidans, has been described. Two isolates of the species differed only in the tendency of one, previously known as strain TH3, to grow in filaments. The chromosomal DNA base composition is between 67 and 69 mol% G + C. The capacity of this species to fix CO2 from air was greater than that of iron-oxidizing thermoacidophiles of the genus Sulfobacillus, which required an enhanced CO2 concentration for optimum autotrophic growth. Under air, ferrous iron oxidation in mixed cultures of A. ferrooxidans with either Sulfobacillus thermosulfidooxidans or Sulfobacillus acidophilus was more extensive than in pure cultures of these three strains. The greater part of ferrous iron oxidation in mixed cultures probably resulted from activity of the Sulfobacillus species, which possess a greater tolerance of ferric iron, and which presumably grew mixotrophically utilizing organic compounds from A. ferrooxidans.

Characteristics of Sulfobacillus acidophilus sp. nov. and other moderately thermophilic mineral-sulphide-oxidizing bacteria.

Several isolates of Gram-positive, acidophilic, moderately thermophilic, ferrous-iron- and mineral-sulphide-oxidizing bacteria were examined to establish unequivocally the characteristics of Sulfobacillus-like bacteria. Two species were evident: Sulfobacillus thermosulfidooxidans with 48-50 mol% G+C and Sulfobacillus acidophilus sp. nov. with 55-57 mol% G+C. Both species grew autotrophically and mixotrophically on ferrous iron, on elemental sulphur in the presence of yeast extract, and heterotrophically on yeast extract. Autotrophic growth on sulphur was consistently obtained only with S. acidophilus.