Individual life histories: neither slow nor fast, just diverse.

The slow-fast continuum is a commonly used framework to describe variation in life-history strategies across species. Individual life histories have also been assumed to follow a similar pattern, especially in the pace-of-life syndrome literature. However, whether a slow-fast continuum commonly explains life-history variation among individuals within a population remains unclear. Here, we formally tested for the presence of a slow-fast continuum of life histories both within populations and across species using detailed long-term individual-based demographic data for 17 bird and mammal species with markedly different life histories. We estimated adult lifespan, age at first reproduction, annual breeding frequency, and annual fecundity, and identified the main axes of life-history variation using principal component analyses. Across species, we retrieved the slow-fast continuum as the main axis of life-history variation. However, within populations, the patterns of individual life-history variation did not align with a slow-fast continuum in any species. Thus, a continuum ranking individuals from slow to fast living is unlikely to shape individual differences in life histories within populations. Rather, individual life-history variation is likely idiosyncratic across species, potentially because of processes such as stochasticity, density dependence, and individual differences in resource acquisition that affect species differently and generate non-generalizable patterns across species.

Niches of three sympatric montane ground-dwelling squirrels: Relative importance of climate, topography, and landcover.

Species with different ecological niches will likely exhibit distinct responses to a changing environment. Differences in the magnitude of niche specialization may also indicate which species may be more vulnerable to environmental change, as many life-history characteristics are known to affect climate change vulnerability. We characterized the niche space of three sympatric high-elevation ground-dwelling squirrels, yellow-bellied marmot (Marmota flaviventer), Belding's ground squirrel (Urocitellus beldingi), and golden-mantled ground squirrel (Callospermophilus lateralis), in the alpine and upper subalpine regions of the Sierra Nevada in California. We used 5879 observations of individual squirrels, collected from 4 years (2009-2012) of transect survey data, to quantify which ecogeographical variable types (climate, topography, or landcover) were most important in defining the niche of each species. We conducted Ecological Niche Factor Analysis to quantify the niche and generate indices of "marginality" (magnitude of selection) and "specialization" (narrowness of niche space). All three species demonstrated differential use of niche space when compared to the available niche space. Moreover, the relative importance of the variables shaping the niche differed among these species. For example, the presence of meadows was important in defining the niche for U. beldingi and M. flaviventer, but the presence of conifers was important to C. lateralis. Precipitation was important in defining the niche for all three species, positively so for U. beldingi, and negatively for the other two species. The niche breadth of these three species was also positively associated with geographic range size. Mammals in high-elevation mountain systems often are perceived as vulnerable to climate shifts, but our results underscore the importance of also including non-climate-based factors in defining the niche. The overall magnitude of niche selection for all three species was driven by a combination of topographic, climatic, and landcover factors; thus, efforts to forecast areas where these species can persist in the future need to evaluate from more than just a climatic perspective.

Current and lagged climate affects phenology across diverse taxonomic groups.

The timing of life events (phenology) can be influenced by climate. Studies from around the world tell us that climate cues and species' responses can vary greatly. If variation in climate effects on phenology is strong within a single ecosystem, climate change could lead to ecological disruption, but detailed data from diverse taxa within a single ecosystem are rare. We collated first sighting and median activity within a high-elevation environment for plants, insects, birds, mammals and an amphibian across 45 years (1975-2020). We related 10 812 phenological events to climate data to determine the relative importance of climate effects on species' phenologies. We demonstrate significant variation in climate-phenology linkage across taxa in a single ecosystem. Both current and prior climate predicted changes in phenology. Taxa responded to some cues similarly, such as snowmelt date and spring temperatures; other cues affected phenology differently. For example, prior summer precipitation had no effect on most plants, delayed first activity of some insects, but advanced activity of the amphibian, some mammals, and birds. Comparing phenological responses of taxa at a single location, we find that important cues often differ among taxa, suggesting that changes to climate may disrupt synchrony of timing among taxa.

Integrated evidence-based extent of occurrence for North American bison (Bison bison) since 1500 CE and before.

Following the near extinction of bison (Bison bison) from its historic range across North America in the late 19th century, novel bison conservation efforts in the early 20th century catalyzed a popular widespread conservation movement to protect and restore bison among other species and places. Since Allen's initial delineation (1876) of the historic distribution of North American bison, subsequent attempts have been hampered by knowledge gaps about bison distribution and abundance prior to and following colonial arrival and settlement. For the first time, we applied a multidisciplinary approach to assemble a comprehensive, integrated geographic database and meta-analysis of bison occurrence over the last 200,000 years, with particular emphasis on the 450 years before present. We combined paleontology, archaeology, and historical ecology data for our database, which totaled 6438 observations. We derived the observations from existing online databases, published literature, and first-hand exploration journal entries. To illustrate the conservative maximum historical extent of occurrence of bison, we created a concave hull using observations occurring over the last 450 years (n = 3379 observations), which is the broadly accepted historical benchmark at 1500 CE covering 59% of the North American continent. Although this distribution represents a historic extent of occurrence-merely delineating the maximum margins of the near-continental distribution-it does not replace a density-based approach reconstructing potential historical range distributions, which identifies core and marginal ranges. However, we envision the observations contained in this database will contribute to further research in the increasingly evidence-based disciplines of bison ecology, evolution, rewilding, management, and conservation. There are no copyright or proprietary restrictions on these data, and this data paper should be cited when the data are reused.

Habitat selection in a fluctuating ground squirrel population: Density-dependence and fitness consequences.

Investigating individual-based habitat settlement decisions is a central theme in ecology, yet studies that quantify density-dependent habitat selection or tie fitness to resource selection decisions remain rare. We quantified habitat selection in golden-mantled ground squirrels (Callospermophilus lateralis) across two spatial scales (home-range placement, and occurrence within the home range) by using 11 consecutive years of data on individual space use, and we used resource selection functions and multilevel modeling to address how habitat preferences may be influenced by density or linked to fitness outcomes. Squirrels preferred dry meadow over other habitat types (wet meadow, aspen, spruce, and willow) at both spatial scales. Squirrels were more likely to use dry meadow that contained shorter vegetation and vision-enhancing prominences such as rocks ("perches"). The use of dry meadow at each scale was not influenced by changes in density. The use of dry meadow did not lead to increased litter size, pre-hibernation mass, or survival. However, squirrels that experienced a greater number of perches or lower local densities had higher survival rates. Our results suggest that a lack of visual obstruction, probably facilitating detection of predators, drives habitat selection in this system. Surprisingly, squirrels maintained their preference for dry meadow as density increased, and they experienced reduced survival as a result. This work furthers our understanding about the causes and consequences of changes in habitat use, informing wildlife management and conservation.

Maternal survival costs in an asocial mammal.

Maternal characteristics, social dynamics, and environmental factors can all influence reproduction and survival and shape trade-offs that might arise between these components of fitness. Short-lived mammals like the golden-mantled ground squirrel (GMGS; Callospermophilus lateralis) tend to maximize effort toward current reproduction at the expense of survival but may be complicated by other aspects of the species' life history and environment. Here, we use 25 years of data (1995-2020) collected from a population of GMGS at the Rocky Mountain Biological Research Laboratory in Gothic, Colorado, to test the effect of several maternal characteristics (e.g., age, experience, and timing of litter emergence), social context (e.g., litter sex ratio and kin density), and environmental context (e.g., date of bare ground and length of vegetative growing season) on survival of reproductive female GMGS using Cox proportional hazard models. Our results indicated that social dynamics (i.e., density) and environmental conditions (i.e., standardized first day of permanent snow cover and length of growing season) explained significant variation in annual maternal survival, while maternal characteristics did not. A higher density of related breeding females and the total number of females (both related and unrelated to the focal mother) were associated with an increase in the mortality hazard. A later standardized date of the first day of permanent snow cover and a shorter growing season both reduced the maternal mortality hazard. Together, our results suggest that factors extrinsic to the squirrels affect maternal survival and thus may also influence local population growth and dynamics in GMGS and other short-lived, territorial mammal species.

Temporal correlations among demographic parameters are ubiquitous but highly variable across species.

Temporal correlations among demographic parameters can strongly influence population dynamics. Our empirical knowledge, however, is very limited regarding the direction and the magnitude of these correlations and how they vary among demographic parameters and species' life histories. Here, we use long-term demographic data from 15 bird and mammal species with contrasting pace of life to quantify correlation patterns among five key demographic parameters: juvenile and adult survival, reproductive probability, reproductive success and productivity. Correlations among demographic parameters were ubiquitous, more frequently positive than negative, but strongly differed across species. Correlations did not markedly change along the slow-fast continuum of life histories, suggesting that they were more strongly driven by ecological than evolutionary factors. As positive temporal demographic correlations decrease the mean of the long-run population growth rate, the common practice of ignoring temporal correlations in population models could lead to the underestimation of extinction risks in most species.

Social mating system and sex-biased dispersal in mammals and birds: a phylogenetic analysis.

The hypothesis that patterns of sex-biased dispersal are related to social mating system in mammals and birds has gained widespread acceptance over the past 30 years. However, two major complications have obscured the relationship between these two behaviors: 1) dispersal frequency and dispersal distance, which measure different aspects of the dispersal process, have often been confounded, and 2) the relationship between mating system and sex-biased dispersal in these vertebrate groups has not been examined using modern phylogenetic comparative methods. Here, we present a phylogenetic analysis of the relationship between mating system and sex-biased dispersal in mammals and birds. Results indicate that the evolution of female-biased dispersal in mammals may be more likely on monogamous branches of the phylogeny, and that females may disperse farther than males in socially monogamous mammalian species. However, we found no support for a relationship between social mating system and sex-biased dispersal in birds when the effects of phylogeny are taken into consideration. We caution that although there are larger-scale behavioral differences in mating system and sex-biased dispersal between mammals and birds, mating system and sex-biased dispersal are far from perfectly associated within these taxa.

Stochastic population dynamics of a montane ground-dwelling squirrel.

Understanding the causes and consequences of population fluctuations is a central goal of ecology. We used demographic data from a long-term (1990-2008) study and matrix population models to investigate factors and processes influencing the dynamics and persistence of a golden-mantled ground squirrel (Callospermophilus lateralis) population, inhabiting a dynamic subalpine habitat in Colorado, USA. The overall deterministic population growth rate λ was 0.94±SE 0.05 but it varied widely over time, ranging from 0.45±0.09 in 2006 to 1.50±0.12 in 2003, and was below replacement (λ<1) for 9 out of 18 years. The stochastic population growth rate λ(s) was 0.92, suggesting a declining population; however, the 95% CI on λ(s) included 1.0 (0.52-1.60). Stochastic elasticity analysis showed that survival of adult females, followed by survival of juvenile females and litter size, were potentially the most influential vital rates; analysis of life table response experiments revealed that the same three life history variables made the largest contributions to year-to year changes in λ. Population viability analysis revealed that, when the influences of density dependence and immigration were not considered, the population had a high (close to 1.0 in 50 years) probability of extinction. However, probability of extinction declined to as low as zero when density dependence and immigration were considered. Destabilizing effects of stochastic forces were counteracted by regulating effects of density dependence and rescue effects of immigration, which allowed our study population to bounce back from low densities and prevented extinction. These results suggest that dynamics and persistence of our study population are determined synergistically by density-dependence, stochastic forces, and immigration.

Proximate causes of natal dispersal in female yellow-bellied marmots, Marmota flaviventris.

We investigated factors influencing natal dispersal in 231 female yearling yellow-bellied marmots (Marmota flaviventris) using comprehensive analysis of 10 years (1983-1993) of radiotelemetry and 37 years (1963-1999) of capture-mark-recapture data. Only individuals whose dispersal status was verified, primarily by radiotelemetry, were considered. Univariate analyses revealed that six of the 24 variables we studied significantly influenced dispersal: dispersal was less likely when the mother was present, amicable behavior with the mother and play behavior were more frequent, and spatial overlap was greater with the mother, with matriline females, and with other yearling females. Using both univariate and multivariate analyses, we tested several hypotheses proposed as proximate causes of dispersal. We rejected inbreeding avoidance, population density, body size, social intolerance, and kin competition as factors influencing dispersal. Instead, our results indicate that kin cooperation, expressed via cohesive behaviors and with a focus on the mother, influenced dispersal by promoting philopatry. Kin cooperation may be an underappreciated factor influencing dispersal in both social and nonsocial species.

Influence of local demography on asymptotic and transient dynamics of a yellow-bellied marmot metapopulation.

Despite recent advances in biodemography and metapopulation ecology, we still have limited understanding of how local demographic parameters influence short- and long-term metapopulation dynamics. We used long-term data from 17 local populations, along with the recently developed methods of matrix metapopulation modeling and transient sensitivity analysis, to investigate the influence of local demography on long-term (asymptotic) versus short-term (transient) dynamics of a yellow-bellied marmot metapopulation in Colorado. Both long- and short-term dynamics depended primarily on a few colony sites and were highly sensitive to changes in demography at these sites, particularly in survival of reproductive adult females. Interestingly, the relative importance of sites differed between long- and short-term dynamics; the spatial structure and local population sizes, while insignificant for asymptotic dynamics, were influential on transient dynamics. However, considering the spatial structure was uninformative about the relative influence of local demography on metapopulation dynamics. The vital rates that were the most influential on local dynamics were also the most influential on both long- and short-term metapopulation dynamics. Our results show that an explicit consideration of local demography is essential for a complete understanding of the dynamics and persistence of spatially structured populations.

Home range and parasite diversity in mammals.

Parasite diversity among and within host species is not solely the result of random processes; rather, it depends on a suite of physiological or ecological host traits as well as environmental factors. Because most macroparasites exhibit life cycles that include infective stages off the definitive host and that rely on host movements for dissemination, parasite acquisition by a host depends largely on hosts being present in a given area where and when infective stages are present. Consequently, host ranging pattern may have a major influence on parasite diversity. Larger home range size is hypothesized to be associated with higher parasite species richness because hosts living in large home ranges should encounter a greater diversity of habitats and other host individuals, which in turn may favor infection by a great diversity of parasite species. By focusing on helminths in wild mammals, we show that an increase in home range area does not lead to an increase in parasite diversity in ungulates and, moreover, that it is associated with a decrease in parasite species richness in carnivores and in glires (rodents and lagomorphs). We also show that home range size is negatively correlated with host density in mammals after correcting both variables for host body mass. We discuss these results from an epidemiological perspective.

Threat of hantavirus pulmonary syndrome to field biologists working with small mammals.

Low risk for hantavirus pulmonary syndrome (HPS) has been reported among biologists engaged in fieldwork with rodents. The overall probability of acquiring HPS when working with rodents appears to be 1 in 1,412 (0.00071). Nonetheless, a causal link between HPS and lack of personal protective equipment (PPE) use is suggested by some investigators. However, supporting data are incomplete and consequently misleading. A recent HPS case was assumed to be acquired during rodent-handling activities, although substantial peridomestic exposure was evident. Regulatory groups interpret inadequate data as evidence of the need for excessive and inappropriate PPE, which can hamper field research and instructional efforts. PPE recommendations should be reviewed and revised to match the risk associated with different types of fieldwork with small mammals.

Hematology and serum chemistry of the island spotted skunk on Santa Cruz Island.

We determined serum biochemistry and hematologic values for island spotted skunks (Spilogale gracilis amphiala) on Santa Cruz Island (California, USA). Samples were collected from island spotted skunks chemically restrained with ketamine hydrochloride and acepromazine in August 1999 (dry season) and from skunks manually restrained in August 2000 (dry season) and January 2001 (wet season). One parameter, glucose, significantly differed with season, with higher levels during the wet season. Serum chemistry and hematologic profiles suggest that method of restraint (manual or chemical), as well as other methodologic details, may influence blood characteristics in the island spotted skunk.