Effect of energy availability, seasonality, and geographic range on brown bear life history

Life-history theory allows predictions of how changes in environmental selection pressures along a species' geographic distribution result in discrete shifts in life-history traits. We tested for spatial patterns of 24 populations of brown bears Ursus arctos across North America that grouped according to the following environmental and population parameters: evapotranspiration as a correlate of primary productivity of vegetation, coefficient of variation of monthly evapotranspiration values as a measure of seasonality. population density, and adult female weight. Cluster analysis grouped brown bear populations into two regions: Pacific-coastal populations characterized by high population density and large females that lived in areas of high primary productivity and low seasonality. and inland and barren-ground populations characterized by relatively low density and small bears that lived in areas of low productivity and high seasonality. For each region, we tested whether life-history traits (age at maturity and interbirth interval) related to primary productivity or seasonality. High altitude (interior: > 1000 m) and high latitude (barren-ground; >65°N) populations respond to extremes in seasonality with risk-spreading adaptations. For example, age at maturity and interbirth interval increased with greater seasonality. In contrast, Pacific-coastal populations living on the western edge of brown bear geographic range respond to intraspecific competition at high densities by maximizing offspring competitive ability. For example, age at maturity increased with greater primary productivity and high population density. In each region, the female parent decided on the life-history trade-offs required to reduce the risks of offspring mortality depending on the environmental pattern.     

Length‐weight relationship and biological data of a threatened fish,Ptychobarbus chungtienensis (Tsao, 1964) in Bita Lake of Shangri‐La,Yunnan, China

This paper provides the length‐weight relationship (LWR) and other biological information of a threatened plateau freshwater fish species Ptychobarbus chungtienensis in the subfamily Schizothoracinae of the family Cyprinidae, from the Bita Lake of Shangri‐la, in Yunnan, China. The LWR was BW = 0.00954 TL^2.95 for a total of 392 individuals, where the coefficient b was close to 3.0, suggesting a near‐isometric growth in this species. The total length: standard length relationship was TL = 0.399 + 1.151 SL. Fitting a von Bertalanffy growth function to the observed length‐at‐age data resulted in L_(t) = 53.1 (1‐e^?0.08(t?0.2)). The natural mortality rate (M) was 0.20 year^?1. Length at first maturity (Lm) was 20.57 cm, and 6.3 years the theoretical estimated age at first maturity (Am). Ptychobarbus chungtienensis mature gradually beginning in May, reaching a reproduction peak around August, with most individuals completing reproduction by October. The gonadosomatic index (GSI) was highest in August (GSI = 0.1351) and lowest in October (GSI = 0.0467). The male index (GSI = 0.0866) was lower than in females (GSI = 0.1466) (P < 0.05). Mean condition factor (Km) of P. chungtienensis was 0.8500, and not significantly different between sexes (P = 0.124), but differing significantly between seasons. In October the condition factor (Km = 0.9211) was significantly higher than in May (Km = 0.8379) or in August (Km = 0.8412) (P < 0.05).     

Moose density and habitat productivity affects reproduction,growth and species composition in field layer vegetation

Question: What is the effect of a gradient in moose density on reproduction, growth and functional composition of the field layer vegetation in a boreal forest, and how is this effect modified by habitat productivity? Location: Northwest of Umeå, Västerbotten, northern Sweden. Methods: Field layer vegetation was surveyed in an experimental setup with simulation of three different moose densities and a control in eight study sites along a gradient of habitat productivity. Results: We found that increased moose density led to decreased cover and reproductive effort of a browsed dwarf shrub (bilberry, Vaccinium myrtillus L.) and increased cover and reproductive effort of a non‐browsed graminoid (wavy hair‐grass, Avenella flexuosa (L.) Drejer). Increased moose density led to increased light availability and probably reduced competition from V. myrtillus. Total reproductive effort in the field layer vegetation increased, height decreased and cover of light‐demanding species and graminoids increased with increasing moose density. The effects of moose density were modified by the productivity gradient, leading to a higher relative increase in light availability and reproductive effort in highly productive areas than in low productive areas. Conclusions: Increased light availability was an important indirect effect of moose density, leading to less competition for light and a shift towards early successional species. The effect of moose density on light availability was modified by habitat productivity, leading to stronger relative effects in highly productive areas than in low productive areas.     

Disturbance-Mediated Apparent Competition Decouples in a Northern Boreal Caribou Range

The most widely reported threat to boreal and mountain populations of woodland caribou (Rangifer tarandus caribou; caribou) involves habitat- or disturbance-mediated apparent competition (DMAC). With DMAC, natural and anthropogenic disturbances that increase the abundance of deciduous-browsing cervids (e.g., moose [Alces alces], deer [Odocoileus spp.]) are thought to promote predator (especially wolf [Canis lupus]) numbers, which heightens predation risk to caribou. We know most about the effects of DMAC on caribou where the species is under threat by anthropogenic activities in relatively productive southern boreal and mountain systems. Yet, >60% of extant boreal caribou range in North America consists of northern shield and taiga ecoregions of low productivity where caribou may compete with only 1 ungulate species (moose) in the context of DMAC. In this environment, we know very little of how DMAC acts as a limiting factor to caribou. In Saskatchewan, Canada, from 2014–2018, using a combination of vegetation sampling, aerial surveys, and telemetry data (n = 38 wolves), we searched for evidence of DMAC (trends in data consistent with the hypothesis) in an 87,193-km² section of the Western Boreal Shield, a poorly productive but pristine region (0.18% of land cover classed as an anthropogenic feature) with a historically high fire-return interval (47% of stands aged <40 years). Despite the high levels of disturbance, moose density was relatively low (47 moose/1,000 km²), likely because of the scarcity of deciduous or mixed-wood stands and low abundance of deciduous browse in the young conifer stands that dominated the landscape. In contrast, boreal caribou density was relatively high for the species (37 caribou/1,000 km²). Wolf density (3.1 wolves/1,000 km²) and pack sizes ( = 4.0 wolves/pack) were low and resident (established) territories were large ( = 4,360 km²; 100% minimum convex polygon). The low density of wolves mirrored the low (standardized) ungulate biomass index (UBI; moose + boreal caribou) of the study area (0.36 UBI/km²). We conclude that wolf and hence caribou populations were not responding in accordance with the outcomes generally predicted by DMAC in our study area because the requisite strong, positive response to fire of deciduous-browse and alternate-prey abundance was lacking. As a limiting factor to caribou, DMAC is likely modulated at a macroecological scale by factors such as net primary productivity, a corollary to the general hypothesis that we advance here (i.e., primary productivity hypothesis of DMAC). We caution against managing for caribou based on the presumption of DMAC where the mechanism does not apply, which may include much of boreal caribou range in the north. © 2020 The Wildlife Society.     

The effects of competitor density on aggressive behaviour and resource defence in a Poeciliid fish

To test the hypothesis that competitor density and time spent at a food resource influences aggressive behaviour in male swordtail, Xiphophorus sp., fish were kept at two densities (low: 18 fish m^?3; high: 54 fish m^?3) and aggressive behaviour was recorded with the time at the food resource used as a covariate. Competitor density is likely to affect the cost:benefit ratio of food defence. When density is low, there should be sufficient food for all individuals and intra‐specific interactions are expected to be rare, while at high densities, increased intra‐specific encounter rates mean that individuals may spend more time defending a resource than utilising it. Food resource defence should occur at intermediate densities. The frequency of aggression, i.e. bites, chases, and display behaviour was significantly positively influenced by time spent at the food resource (Bites: F_1,31 = 8.186, P = 0.007; Chases: F_1,31 = 6.439, P = 0.016; Displays: F_1,31 = 4.435, P = 0.043) suggesting that food resource defence occurred. Competitor density had no effect on food resource defence and minimal effect on the frequency of aggression with only one type of aggressive behaviour, male–male displays, showing a difference between densities (F_1,31 = 6.975, P = 0.013). This finding is suggested to be a result of the formation of dominance hierarchies in this species. Aggression from the dominant individual may be directed at only subordinates of the next dominance rank and subordinate behaviour may be restricted by status rather than immediate threat. In such a situation, aggression may be independent of competitor density.     

Mitochondrial DNA phylogeography of Spodoptera exigua across a broad geographic area in China

The beet armyworm, Spodoptera exigua (Lepidoptera: Noctuidae), is an important agriculture pest in China that causes serious economic losses in some of the main crop‐producing areas. To monitor and manage this pest effectively, it is necessary to investigate its phylogeographic patterns in China. In this study, we used a partial sequence of mitochondrial DNA Cytb gene consisting of 724 bps to investigate the genetic diversity of the beet armyworm. A total of 765 individuals from 47 populations across the main distribution range of the species were collected, and 112 haplotypes were identified. Moderate‐to‐high levels of genetic diversity (Hd = 0.672 ± 0.017, Pi = 0.00268 ± 0.00021) for the total populations were obtained. Phylogenetic and median‐joining network analyses indicated there was no distinct geographic distribution pattern among haplotypes. Overall, the study also revealed significant differentiation among some populations (P < 0.05). The F_ST values of Shenyang population (SY2012–SY 2014), as well as Baoding (BD), Taian (TA), Lucheng (LC), Zhengzhou (ZZ) and Wuhan (WH), were significantly different from those of the populations in most other locations. Hierarchical AMOVA showed there was no significantly genetic structure between populations located in seven geographic regions and four main bioclimatic zones. Finally, unimodal mismatch distribution combined with negative Tajima's D (D = ?2.696, P < 0.001) and Fu's F_S (F_S = ?207.228, P > 0.05) indicated recent population expansion of S. exigua at large spatial scales in China.     

Demographic and environmental influences on life-history traits of isolated populations of the Andean catfish Astroblepus ubidiai

In this study, we examined the influence of demographic and environmental variables on the life histories of six remnant populations of the Andean catfishAstroblepus ubidiai (Actinopterygii; Siluriform) located in isolated refuges, and tested six predictions on the relationships between age at maturity, generation time, population density and juvenile and adult growth and survival rates. The three populations inhabiting the watershed with a direct connection to a major lake (Imbakucha) exhibited later maturity, a longer generation time, higher adult survivorship, a higher adult-to-juvenile survival ratio and lower biomass density than those of the adjacent, non-lacustrine watersheds. Across all six populations, there was a strong correlation between mean age at maturity and both the adult-to-juvenile survival ratio and biomass density. Study populations also showed a pattern of inverse density-dependence associated with Allee effects. We conclude that without limitations in the availability of environmental resources, higher population density can be negatively related to age at maturity. This relationship would be expected to become positive when carrying capacity of the systems is reached. Delayed maturity in the Imbakucha populations may be an adaptive response for increasing juvenile survival under stressful conditions.     

Genome-wide SNP analysis of three moose subspecies at the southern range limit in the contiguous United States

Genome-wide evaluations of genetic diversity and population structure are important for informing management and conservation of trailing-edge populations. North American moose (Alces alces) are declining along portions of the southern edge of their range due to disease, species interactions, and marginal habitat, all of which may be exacerbated by climate change. We employed a genotyping by sequencing (GBS) approach in an effort to collect baseline information on the genetic variation of moose inhabiting the species’ southern range periphery in the contiguous United States. We identified 1920 single nucleotide polymorphisms (SNPs) from 155 moose representing three subspecies from five states: A. a. americana (New Hampshire), A. a. andersoni (Minnesota), and A. a. shirasi (Idaho, Montana, and Wyoming). Molecular analyses supported three geographically isolated clusters, congruent with currently recognized subspecies. Additionally, while moderately low genetic diversity was observed, there was little evidence of inbreeding. Results also indicated?>?20% shared ancestry proportions between A. a. shirasi samples from northern Montana and A. a. andersoni samples from Minnesota, indicating a putative hybrid zone warranting further investigation. GBS has proven to be a simple and effective method for genome-wide SNP discovery in moose and provides robust data for informing herd management and conservation priorities. With increasing disease, predation, and climate related pressure on range edge moose populations in the United States, the use of SNP data to identify gene flow between subspecies may prove a powerful tool for moose management and recovery, particularly if hybrid moose are more able to adapt.

     

Evolution of indefinite generation lengths

Intrinsic mortality at relatively advanced age results from natural selection favouring early reproduction. When individuals produce many and early offspring, costs are incurred due to the accumulation of pleiotropic mutations with adverse effects late in life, which make senescence an inevitable result of evolution. However, a few organisms exist with seemingly indefinite generation lengths. This paper identifies conditions of life history and ecology under which natural selection favours continually reducing intrinsic mortality. The analysis considers the particular case of populations held at carrying capacity by density dependence suppressing net fertility, acting either on fecundity equally across fecund ages or on juvenile mortality up to the first fecund age. This broad category of density dependence increases the susceptibility of populations to invasion by mutations that extend net fertility into later life. Whether a population then evolves longer generation times depends on the relative timing of period benefit to onset of senescent cost amongst antagonistic pleiotropisms that arise to confront the selection process. Simulations show that if the onset of senescent costs either precedes the benefit, is concurrent with it, or is postponed by a fixed interval, then natural selection will favour those pleiotropisms that increase generation length. The selection on generation-lengthening pleiotropisms continues indefinitely and regardless of current generation length, for as long as the environment remains constant. The wide range of life histories encompassed by these conditions leads to the conclusion that many species may have evolved generation lengths considerably longer than the minimum set by developmental constraints.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 269–280.     

Components of breeding performance in two competing species: habitat heterogeneity, individual quality and density-dependence

Density‐dependent breeding performance due to habitat heterogeneity has been shown to regulate populations of territorial species, since the progressive occupation of low quality territories as breeding density increases may cause a decline in the mean per capita fecundity of a population while variation in fecundity increases. Although the preemptive use of sites may relegate low quality individuals to sites of progressively lower suitability, few studies on density dependence have tried to separate the effects of territory quality from individual quality, and none have simultaneously considered the effects of heterospecific competitors. Using two long‐term monitored populations, we assessed the relative contribution of habitat heterogeneity and bird quality (in terms of age) on the productivity of sympatric golden Aquila chrysaetos and Bonelli's eagles Hieraaetus fasciatus under different scenarios of intra‐ and inter‐specific competition. Productivity (number of offspring fledged) varied among territories and average annual productivity was negatively related to its variability in both species and populations, thus giving some support to the habitat heterogeneity hypothesis. However, the effect of habitat heterogeneity on productivity became non‐significant when parental age and local density estimators were included in multivariate analyses. Therefore, temporal changes in bird quality (age) combined with intra‐ and interspecific competition explained variability in territory productivity rather than habitat heterogeneity among territories per se. The recruitment of subadult breeders, a surrogate of mortality in eagles, strongly varied among territories. Habitat heterogeneity in productivity may thus arise not because sites differ in suitability for reproduction but because of differences in factors affecting survival. Territories associated with high mortality risks have a higher probability of being occupied by young birds, whose lower quality, interacting with the density competitors, leads to a reduction of productivity. Site‐dependent variability in adult survival and interspecific competition may be extensive, but so far largely overlooked, factors to be seriously considered for the site‐dependent population regulation framework.     

Comparative survival and recovery of Ross's and lesser snow geese from Canada's central arctic

Large increases in several populations of North American arctic geese have resulted in ecosystem-level effects from associated herbivory. Consequently, some breeding populations have shown density dependence in recruitment through declines in food availability. Differences in population trajectories of lesser snow geese (Chen caerulescens caerulescens; hereafter snow geese) and Ross's geese (C. rossii) breeding in mixed-species colonies south of Queen Maud Gulf (QMG), in Canada's central arctic, suggest that density dependence may be limiting snow goose populations. Specifically, long-term declines in age ratios (immature:adult) of harvested snow geese may have resulted from declines in juvenile survival. Thus, we focused on juvenile (first-year) survival of snow and Ross's geese in relation to timing of reproduction (annual mean nest initiation date) and late summer weather. We banded Ross's and snow geese from 1991 to 2008 in the QMG Migratory Bird Sanctuary. We used age-structured mark-recapture models to estimate annual survival rates for adults and juveniles from recoveries of dead birds. Consistent with life history differences, juvenile snow geese survived at rates higher than juvenile Ross's geese. Juvenile survival of both species also was lower in late seasons, but was unrelated to arctic weather measured during a 17-day period after banding. We found no evidence of density dependence (i.e., a decline in juvenile survival over time) in either species. We also found no interspecific differences in age-specific hunting vulnerability, though juveniles were more vulnerable than adults in both species, as expected. Thus, interspecific differences in survival were unrelated to harvest. Lower survival of juvenile Ross's geese may result from natural migration mortality related to smaller body size (e.g., greater susceptibility to inclement weather or predation) compared to juvenile snow geese. Despite lower first-year survival, recruitment by Ross's geese may still be greater than that by snow geese because of earlier sexual maturity, greater breeding propensity, and higher nest success by Ross's geese. © 2012 The Wildlife Society.     

A resurrection study reveals limited evolution of thermal performance in response to recent climate change across the geographic range of the scarlet monkeyflower

Evolutionary rescue can prevent populations from declining under climate change, and should be more likely at high-latitude, “leading” edges of species’ ranges due to greater temperature anomalies and gene flow from warm-adapted populations. Using a resurrection study with seeds collected before and after a 7-year period of record warming, we tested for thermal adaptation in the scarlet monkeyflower Mimulus cardinalis. We grew ancestors and descendants from northern-edge, central, and southern-edge populations across eight temperatures. Despite recent climate anomalies, populations showed limited evolution of thermal performance curves. However, one southern population evolved a narrower thermal performance breadth by 1.31°C, which matches the direction and magnitude of the average decrease in seasonality experienced. Consistent with the climate variability hypothesis, thermal performance breadth increased with temperature seasonality across the species’ geographic range. Inconsistent with performance trade-offs between low and high temperatures across populations, we did not detect a positive relationship between thermal optimum and mean temperature. These findings fail to support the hypothesis that evolutionary response to climate change is greatest at the leading edge, and suggest that the evolution of thermal performance is unlikely to rescue most populations from the detrimental effects of rapidly changing climate.     

Age‐associated microRNA expression in human peripheral blood is associated with all‐cause mortality and age‐related traits

Recent studies provide evidence of correlations of DNA methylation and expression of protein‐coding genes with human aging. The relations of microRNA expression with age and age‐related clinical outcomes have not been characterized thoroughly. We explored associations of age with whole‐blood microRNA expression in 5221 adults and identified 127 microRNAs that were differentially expressed by age at P < 3.3 × 10^?4 (Bonferroni‐corrected). Most microRNAs were underexpressed in older individuals. Integrative analysis of microRNA and mRNA expression revealed changes in age‐associated mRNA expression possibly driven by age‐associated microRNAs in pathways that involve RNA processing, translation, and immune function. We fitted a linear model to predict ‘microRNA age’ that incorporated expression levels of 80 microRNAs. MicroRNA age correlated modestly with predicted age from DNA methylation (r = 0.3) and mRNA expression (r = 0.2), suggesting that microRNA age may complement mRNA and epigenetic age prediction models. We used the difference between microRNA age and chronological age as a biomarker of accelerated aging (Δage) and found that Δage was associated with all‐cause mortality (hazards ratio 1.1 per year difference, P = 4.2 × 10^?5 adjusted for sex and chronological age). Additionally, Δage was associated with coronary heart disease, hypertension, blood pressure, and glucose levels. In conclusion, we constructed a microRNA age prediction model based on whole‐blood microRNA expression profiling. Age‐associated microRNAs and their targets have potential utility to detect accelerated aging and to predict risks for age‐related diseases.     

Challenges and opportunities for estimating abundance of a low-density moose population

Monitoring large herbivores across their core range has been readily accomplished using aerial surveys and traditional distance sampling. But for peripheral populations, where individuals may occur in patchy, low-density populations, precise estimation of population size and trend remains logistically and statistically challenging. For moose (Alces alces) along their southern range margin in northern New York, USA, we sought robust estimates of moose distribution, abundance, and population trend (2016–2019) using a combination of aerial surveys (line transect distance-sampling), repeated surveys in areas where moose were known to occur to boost the number of detections, and density surface modeling (DSM) with spatial covariates. We achieved a precise estimate of density (95% CI = 0.00–0.29 moose/km²) for this small population (656 moose, 95% CI = 501–859), which was patchily distributed across a large and heavily forested region (the 24,280-km² Adirondack Park). Local moose abundance was positively related to active timber management, elevation, and snow cover, and negatively related to large bodies of water. As expected, moose abundance in this peripheral population was low relative to its core range in other northern forest states. Yet, in areas where abundance was greatest, moose densities in New York approached those where epizootics of winter tick (Dermacentor albipictus) have been reported, underscoring the need for effective and efficient monitoring. By incorporating autocorrelation in observations and landscape covariates, DSM provided spatially explicit estimates of moose density with greater precision and no additional field effort over traditional distance sampling. Combined with repeated surveys of areas with known moose occurrence to achieve viable sample sizes, DSM is a useful tool for effectively monitoring low density and patchy populations.     

The Pillars of Hercules as a bathymetric barrier to gene flow promoting isolation in a global deep‐sea shark (Centroscymnus coelolepis)

Knowledge of the mechanisms limiting connectivity and gene flow in deep‐sea ecosystems is scarce, especially for deep‐sea sharks. The Portuguese dogfish (Centroscymnus coelolepis) is a globally distributed and near threatened deep‐sea shark. C. coelolepis population structure was studied using 11 nuclear microsatellite markers and a 497‐bp fragment from the mtDNA control region. High levels of genetic homogeneity across the Atlantic (Φ_ST = ?0.0091, F_ST = 0.0024, P > 0.05) were found suggesting one large population unit at this basin. The low levels of genetic divergence between Atlantic and Australia (Φ_ST = 0.0744, P < 0.01; F_ST = 0.0015, P > 0.05) further suggested that this species may be able to maintain some degree of genetic connectivity even across ocean basins. In contrast, sharks from the Mediterranean Sea exhibited marked genetic differentiation from all other localities studied (Φ_ST = 0.3808, F_ST = 0.1149, P < 0.001). This finding suggests that the shallow depth of the Strait of Gibraltar acts as a barrier to dispersal and that isolation and genetic drift may have had an important role shaping the Mediterranean shark population over time. Analyses of life history traits allowed the direct comparison among regions providing a complete characterization of this shark's populations. Sharks from the Mediterranean had markedly smaller adult body size and size at maturity compared to Atlantic and Pacific individuals. Together, these results suggest the existence of an isolated and unique population of C. coelolepis inhabiting the Mediterranean that most likely became separated from the Atlantic in the late Pleistocene.     

Impact of simulated moose densities on abundance and richness of vegetation,herbivorous and predatory arthropods along a productivity gradient

Large herbivores can affect vegetation structure and species composition as well as material and energy flows in the ecosystem through their selective feeding, defecation, urination and trampling. These changes have a large potential to indirectly affect other trophic levels, but the mechanisms are poorly known. We studied the impacts of moose Alces alces browsing along a gradient of site productivity by experimentally simulating four different moose densities. Here we show that moose can affect the richness and abundance of three trophic levels in Swedish boreal forests through complex direct and indirect impacts, but in qualitatively different ways depending on how the physical habitat or food resources of a trophic level are affected. Vegetation richness had a hump‐shaped (unimodal) response to increased moose density. Leaf litter production decreased when browsing increased, which in turn depressed the abundance of flying prey for spiders. Consequently, spider abundance and richness declined monotonically. The responses of spider richness to moose density were further conditioned by site productivity: the response was positive at productive and negative at unproductive sites. In contrast, herbivorous Hemiptera were not affected by moose, most likely because the abundance of their food plants was not affected. The highest simulated moose density had an impact on all variables responding to moose even after a few years of treatment and can be considered as overabundance. We also show that the impacts of low or moderate moose density can be positive to some of the organisms negatively affected by high density. The level of herbivore population density that leads to substantial community impacts also depends on site factors, such as productivity.     

Energy availability, abundance, energy-use and species richness in forest bird communities: a test of the species–energy theory

Aim To test the ‘more individuals hypothesis’ as a mechanism for the positive association between energy availability and species richness. This hypothesis predicts that total density and energy use in communities is linearly related to energy availability, and that species richness is a positive function of increased density. We also evaluate whether similar energy–density patterns apply to different migratory groups (residents, short‐distance migrants and tropical migrants) separately. Location European and North American forest bird communities. Methods We collected published breeding bird census data from Europe and North America (n = 187). From each census data we calculated bird density (pairs 10 ha^?1), energy use by the community (the sum of metabolic needs of individuals, Watts 10 ha^?1) and geographical location with an accuracy of 0.5°. For each bird census data coordinate we extracted the corresponding monthly values of actual evapotranspiration (AET). From these values we calculated corresponding AET values that we expected to explain the density energy use of forest birds: total annual, breeding season (June) and winter AET. We used general linear modelling to analyse these data controlling for the area of census plots, forest type and census method. Results Total density and energy use in European and North American forest bird communities were linear functions of annual productivity, and increased density and energy use then translated into more species. Also resident bird density and energy consumption were positive functions of annual productivity, but the relationship between productivity and density as well as between productivity and energy use was weaker for migrants. Main conclusions Our results are consistent with the more individuals hypothesis that density and energy use in breeding forest bird communities is coupled tightly with the productivity of the environment, and that increased density and energy consumption results in more species. However, not all community members (migratory groups) are limited by productivity on the breeding grounds.     

Geographical and latitudinal variation in growth patterns and adult body size of Swedish moose (Alces alces)

We examined the geographical pattern in growth and adult body size among 14 populations of Swedish moose (Alces alces) using data from 4,294 moose (1.5 years old) killed during the hunting season in 1989–1992. In both sexes, adult body mass was significantly positively correlated with latitude. Moose in northern populations had a 15–20% larger adult body mass than moose in the south. Juvenile body mass was correlated with neither latitude nor adult body mass. Thus, variation in time (years) and rate of body growth after the juvenile stage were responsible for most of the variation in adult body mass among populations. Moose in northern populations grew for approximately 2 more years of life than southern moose. In contrast to adult body mass, skeletal size (measured as jawbone length) was not correlated with latitude, suggesting that variation in adult body mass was primarily due to differences in fat reserves. Discrimination between population characteristics, such as moose density, climate, and the amount of browse available to moose, showed climatic harshness to be the most important variable explaining geographical variation in body mass among populations. The results support the notion that in mammals body size increases with latitude in accordance with Bergmann's rule. We conclude that (1) variation in patterns of growth after the juvenile stage is the main cause of the latitudinal trend in adult body size in moose, and (2) climatic conditions are a more important factor than population density and availability of food in explaining geographical variation in growth patterns and adult body mass between populations of Swedish moose.     

Lifetime Sex-Specific Moose Mortality During an Intentional Population Reduction

Where elevated harvest of ungulates is a priority, managers benefit by understanding how various sources of mortality affect the age and sex structure and trend of ungulate populations. Prior studies reported a long period (1997–2014) of moose (Alces alces gigas) nutritional stress from overabundance in our study area, an intentional 31% reduction in moose numbers using liberal harvests of females (2004–2012), and low bear (Ursus spp.) predation and high moose harvest densities relative to other largely roadless systems with moose, bears, and wolves (Canis lupus). In this paper, we detailed management findings after describing causes and rates of mortality from 226 female and 164 male moose radio-collared at 9 months of age (1997–2008) and followed through life (1997–2019) and throughout the population reduction. We listened for mortality signals on radio-collars 1–2 times/month when snow cover was complete and 2–4 times/month when snow cover was incomplete. Upon hearing a mortality signal, we investigated mortality sites usually within 24 hours via helicopter. Excluding hunter-caused mortality, we estimated 28% annual mortality for male yearlings versus 17% for female yearlings, then low annual mortality rates (0–4%) to 84 months of age for males and 96 months of age for females, and gradually increasing annual mortality rates thereafter. Most (83%) male moose ≥24 months of age died from hunters; minor causes included wolves (8%), malnutrition or disease (5%), grizzly bears (U. arctos; 2%), and accidents (2%). Most female moose ≥24 months of age died from wolves (37%) or hunters (33%); minor causes included malnutrition or disease (15%), grizzly bears (10%), and accidents (5%). The proportion of radio-collared females killed by hunters varied depending on numbers of permits issued to hunters; the kill rate of females ≥24 months of age was 58% during the initial 4 years of the 9-year reduction, moderated at 29% during the final 5 years of the reduction, and was only 7% for all other study years. We attributed 32% of hunter kills to illegal harvest and unrecovered hunter kills. Hunters played a key role in the intentional population reduction by harvesting prime-age and near prime-age male and female moose that rarely died from other sources of mortality compared with calf, yearling, and older moose. Restricting general season hunters to primarily harvesting prime-age and older male moose with antler spreads ≥127 cm did not appreciably reduce harvest of adult males. Male moose 2.0–5.3 years of age rarely died from non-hunter causes and were largely harvested at older, prime ages (5.3–8.3 yr of age). Yearling moose of both sexes died primarily from wolves, with wolves selecting more for males. By using liberal harvests of female moose to reduce the population, managers improved moose nutrition and reproduction, met mandates for elevated harvests, and may have avoided a reoccurrence of a previous precipitous decline in moose numbers that was initiated by overabundance and extreme snow depths. © 2019 The Wildlife Society.     

Signals of large scale climate drivers,hatchery enhancement,and marine factors in Yukon River Chinook salmon survival revealed with a Bayesian life history model

Understanding how species might respond to climate change involves disentangling the influence of co‐occurring environmental factors on population dynamics, and is especially problematic for migratory species like Pacific salmon that move between ecosystems. To date, debate surrounding the causes of recent declines in Yukon River Chinook salmon (Oncorhynchus tshawytscha) abundance has centered on whether factors in freshwater or marine environments control variation in survival, and how these populations at the northern extremity of the species range will respond to climate change. To estimate the effect of factors in marine and freshwater environments on Chinook salmon survival, we constructed a stage‐structured assessment model that incorporates the best available data, estimates incidental marine bycatch mortality in trawl fisheries, and uses Bayesian model selection methods to quantify support for alternative hypotheses. Models fitted to two index populations of Yukon River Chinook salmon indicate that processes in the nearshore and marine environments are the most important determinants of survival. Specifically, survival declines when ice leaves the Yukon River later in the spring, increases with wintertime temperature in the Bering Sea, and declines with the abundance of globally enhanced salmon species consistent with competition at sea. In addition, we found support for density‐dependent survival limitations in freshwater but not marine portions of the life cycle, increasing average survival with ocean age, and age‐specific selectivity of bycatch mortality in the Bering Sea. This study underscores the utility of flexible estimation models capable of fitting multiple data types and evaluating mortality from both natural and anthropogenic sources in multiple habitats. Overall, these analyses suggest that mortality at sea is the primary driver of population dynamics, yet under warming climate Chinook salmon populations at the northern extent of the species’ range may be expected to fare better than southern populations, but are influenced by foreign salmon production.