The peppered moth and industrial melanism: evolution of a natural selection case study

From the outset multiple causes have been suggested for changes in melanic gene frequency in the peppered moth Biston betularia and other industrial melanic moths. These have included higher intrinsic fitness of melanic forms and selective predation for camouflage. The possible existence and origin of heterozygote advantage has been debated. From the 1950s, as a result of experimental evidence, selective predation became the favoured explanation and is undoubtedly the major factor driving the frequency change. However, modelling and monitoring of declining melanic frequencies since the 1970s indicate either that migration rates are much higher than existing direct estimates suggested or else, or in addition, non-visual selection has a role. Recent molecular work on genetics has revealed that the melanic (carbonaria) allele had a single origin in Britain, and that the locus is orthologous to a major wing patterning locus in Heliconius butterflies. New methods of analysis should supply further information on the melanic system and on migration that will complete our understanding of this important example of rapid evolution.     

A bird’s eye view of the peppered moth

Industrial melanism in Biston betularia is one of the best known examples of the role of natural selection in evolution and has received considerable scrutiny for many years. The rise in frequency of the dark form of the moth (carbonaria) and a decrease in the pale form (typica) was the result of differential predation by birds, the melanic form being more cyptic than typica in industrial areas where the tree bark was darkened by air pollution. One important aspect of early work evaluating the relative crypsis of the forms of B. betularia on tree trunks with different lichen flora was the reliance on human observers. Humans, however, do not have the same visual capabilities as birds. Birds have well‐developed ultraviolet (UV) vision, an important component of their colour processing system that affects many aspects of behaviour, including prey detection. We examined the UV characteristics of the two forms of B. betularia and a number of foliose and crustose lichens. In human visible light the speckled form typica appeared cyptic when seen against a background of foliose lichen, whereas the dark form carbonaria was conspicuous. Under UV light the situation was reversed. The foliose lichens absorbed UV and appeared dark as did carbonaria. Typica, however, reflected UV and was conspicuous. Against crustose lichens, typica was less visible than carbonaria in both visible and UV light. These findings are considered in relation to the distribution and recolonization of trees by lichens and the resting behaviour of B. betularia.     

Post-biological control invasion trajectory for Melaleuca quinquenervia in a seasonally inundated wetland

The recruitment and mortality of Melaleuca quinquenervia seedlings were evaluated over a 3-year period in a seasonally inundated wetland in the western Everglades region. The mean (±SE) density of seedlings/saplings m^?1 declined from 64.8 (±4.5) to 0.5 (±0.2) over the 3 years, a population reduction of 99.2%. Four distinct water regimes characterized this site: dry, dry to wet transition, flooded, and wet to dry transition. Seedling recruitment was highest in the dry to wet transition and lowest in the flooded water regime, while mortality was highest under flooded and dry water regimes. The mean estimate of population growth (λ) across water regimes was 0.64 ± 0.05 indicating negative population growth. Elimination of introduced insect herbivores using insecticides did not reduce mortality of recruited M. quinquenervia seedlings/saplings indicating that direct herbivory was not responsible for the decline in seedling density. On the other hand, a mean of only 0.2 (±0.03) viable seeds m^?2 d^?1 fell into the plots, an amount considerably lower than in previous studies. We submit that change in the invasion trajectory M. quinquenervia was most likely caused by reduced seed inputs from aerial seed banks depleted by insect herbivory rather than direct herbivory on seedlings. This may indicate a fundamental alteration of M. quinquenervia population dynamics ultimately resulting in a less invasive and, therefore, less ecologically damaging species.     

Intersubtype Differences in the Effect of a Rare p24 Gag Mutation on HIV-1 Replicative Fitness

Certain immune-driven mutations in HIV-1, such as those arising in p24^Gag, decrease viral replicative capacity. However, the intersubtype differences in the replicative consequences of such mutations have not been explored. In HIV-1 subtype B, the p24^Gag M250I mutation is a rare variant (0.6%) that is enriched among elite controllers (7.2%) (P = 0.0005) and appears to be a rare escape variant selected by HLA-B58 supertype alleles (P < 0.01). In contrast, in subtype C, it is a relatively common minor polymorphic variant (10 to 15%) whose appearance is not associated with a particular HLA allele. Using site-directed mutant viruses, we demonstrate that M250I reduces in vitro viral replicative capacity in both subtype B and subtype C sequences. However, whereas in subtype C downstream compensatory mutations at p24^Gag codons 252 and 260 reduce the adverse effects of M250I, fitness costs in subtype B appear difficult to restore. Indeed, patient-derived subtype B sequences harboring M250I exhibited in vitro replicative defects, while those from subtype C did not. The structural implications of M250I were predicted by protein modeling to be greater in subtype B versus C, providing a potential explanation for its lower frequency and enhanced replicative defects in subtype B. In addition to accounting for genetic differences between HIV-1 subtypes, the design of cytotoxic-T-lymphocyte-based vaccines may need to account for differential effects of host-driven viral evolution on viral fitness.     

Queen mating frequency and maternity of males in the stingless bee Trigona carbonaria Smith

Summary. Aspects of the sociobiology of the native Australian stingless bee Trigona carbonaria (Hymenoptera, Meliponini) were investigated using highly variable microsatellite markers. The queen mating frequency was estimated by examining genotypes of samples of workers from five colonies across three to five microsatellite loci. In each case, results were consistent with the workers being progeny of a queen mated with a single male. Microsatellite analysis of haploid males from two of the colonies suggested they predominantly arose from queen-laid eggs. As workers are more related to sons-of-workers than to sons-of-queens in monandrous colonies, this is somewhat surprising, and suggests that there may be colony-level costs associated with worker reproduction, or that queens are able to regulate worker reproduction via the ritualized cell provisioning and oviposition process. Males from another T. carbonaria colony were found to be diploid.     

Human-resource subsidies alter the dietary preferences of a mammalian top predator

Seasonal fitness declines are common, but the relative contribution of different reproductive components to the seasonal change in the production of reproductive young, and the component-specific drivers of this change is generally poorly known. We used long-term data (17 years) on breeding time (i.e. date of first egg laid) in northern wheatears (Oenanthe oenanthe) to investigate seasonal reproductive patterns and estimate the relative contributions of reproductive components to the overall decline in reproduction, while accounting for factors potentially linked to seasonal declines, i.e. individual and habitat quality. All reproductive components—nest success (reflecting nest predation rate), clutch size, fledging success and recruitment success—showed a clear decline with breeding time whereas subsequent adult survival did not. A non-linear increase in nest predation rate caused nest success to decline rapidly early in the season and level off at ~80 % success late in the breeding season. The combined seasonal decline in all reproductive components caused the mean production of recruits per nest to drop from around 0.7–0.2; with the relative contribution greatest for recruitment success which accounted for ~50 % of the decline. Our data suggest that changing environmental conditions together with effects of nest predation have strong effects on the seasonal decline in fitness. Our demonstration of the combined effects of all reproductive components and their relative contribution shows that omitting data from later stages of breeding (recruitment) can greatly underestimate seasonal fitness declines.     

Experimental evolution of bet hedging under manipulated environmental uncertainty in Neurospora crassa

All organisms are faced with environmental uncertainty. Bet-hedging theory expects unpredictable selection to result in the evolution of traits that maximize the geometric-mean fitness even though such traits appear to be detrimental over the shorter term. Despite the centrality of fitness measures to evolutionary analysis, no direct test of the geometric-mean fitness principle exists. Here, we directly distinguish between predictions of competing fitness maximization principles by testing Cohen''s 1966 classic bet-hedging model using the fungus Neurospora crassa. The simple prediction is that propagule dormancy will evolve in proportion to the frequency of ‘bad’ years, whereas the prediction of the alternative arithmetic-mean principle is the evolution of zero dormancy as long as the expectation of a bad year is less than 0.5. Ascospore dormancy fraction in N. crassa was allowed to evolve under five experimental selection regimes that differed in the frequency of unpredictable ‘bad years’. Results were consistent with bet-hedging theory: final dormancy fraction in 12 genetic lineages across 88 independently evolving samples was proportional to the frequency of bad years, and evolved both upwards and downwards as predicted from a range of starting dormancy fractions. These findings suggest that selection results in adaptation to variable rather than to expected environments.     

Frequency-Dependent Selection at the PGM-1 Locus of DROSOPHILA PSEUDOOBSCURA

Frequency-dependent fitness was studied at the Pgm-1 locus of Drosophila pseudoobscura with respect to two fitness components: rate of development and larva-to-adult survival. The Pgm-1 locus is very polymorphic with only two alleles, Pgm-1¹⁰⁰ and Pgm-1¹⁰⁴, occurring at high frequencies. For each of these two alleles, 20 homozygous strains were obtained from a sample of 1,140 wild-inseminated females. First-instar larvae of the two genotypes were combined in a set of eight different frequencies: 0.0, 0.10, 0.25, 0.40, 0.60, 0.75, 0.90, and 1.0. Frequency-dependent fitness effects were observed for the two survival-related fitness components examined: larvae of the less common genotype develop faster and have a higher probability of survival than larvae of the more common genotype. The rate of survival at intermediate genotypic frequencies is similar to that in pure cultures. If selection acted solely as frequency-dependent effects on survival-related components of fitness, the equilibrium frequency of the Pgm-1¹⁰⁰ allele would be 0.615 for a two-genotype system, which fits an observed frequency range for this allele in nature between 0.55 and 0.71. Experimentally created linkage disequilibrium was excluded from the experiment by using a large number of independent strains. It is nevertheless possible that the frequency-dependent selection may not affect the Pgm-1 locus per se, but may reflect a linkage disequilibrium present in the natural population. Even if this were the case, the frequency-dependent selection could affect the frequency of the Pgm-1 alleles in nature.     

Impact of Striped-Squirrel Nectar-Robbing Behaviour on Gender Fitness in Alpinia roxburghii Sweet (Zingiberaceae)

Nectar-robbing has the potential to strongly affect male and female reproductive fitness of plants. One example of nectar theft is that shown by striped-squirrels (Tamiops swinhoei) on a number of ginger species, including Alpinia roxburghii and A. kwangsiensis (Zingiberaceae). In this study, we used a fluorescent dye as a pollen analogue, and measured fruit and seed output, to test the effect of squirrel nectar-robbing on A. roxburghii reproductive fitness. Pollen transfer between robbed and unrobbed flowers was assessed by comparing 60 randomly established plots containing robbed and unrobbed flowers. The frequency of squirrel robbing visits and broken styles were recorded from a number of flowers for five consecutive days. Two bee species (Bombus eximius and Apis cerana), were the primary pollinators, and their visitation frequency was recorded for six consecutive days. The results showed that fluorescent powder from unrobbed flowers was dispersed further, and to a greater number of flowers than that placed on robbed flowers. Additionally, robbing flowers caused significant damage to reproductive organs, resulting in lower fruit and seed sets in robbed than in unrobbed flowers and influencing both male and female fitness. The frequency of the primary pollinator visits (B. eximius) was significantly higher for unrobbed plants than for robbed plants. The present study clearly shows the negative impact of squirrel robbing on A. roxburghii male reproductive fitness and neutral impact on female reproductive fitness.     

Density- and Frequency-Dependent Selection at the Mdh-2 Locus in DROSOPHILA PSEUDOOBSCURA

We have studied differences in the number of Drosophila pseudoobscura produced in a culture when the flies differ with respect to two alleles (F and S) at the Mdh-2 locus, which codes for a malate dehydrogenase enzyme. The studies were done at low and at high density in two- and three-genotype combinations (S/S, F/F and S/F), with one-genotype cultures as controls.——Density affects the fitness of the Mdh-2 genotypes. Different genotypes are differently affected, and the genotype of the competitors also makes a difference on the fitness of a given genotype. When three genotypes are present in a culture, particularly at high density, intergenotypic competition is less intense than intragenotypic competition at several frequency combinations. That is, there is "overcompensation": the three genotypes together exploit the environmental resources better than one genotype alone.—The fitness of the genotypes is frequency dependent in both two-genotype and three-genotype combinations. An inverse relationship between frequency and fitness is observed at high density. This may lead to a stable polymorphism, because the fitness of a genotype increases as its frequency decreases.—Forty independent strains, sampled from a natural population, were used in the experiments. This ensures that more than 95% of the variation present in the genome in the natural population is also present is the experimental cultures. It also ensures that the genetic background of the Mdh-2 alleles is randomized in the same way as it is in nature. However, the possibility remains that Mdh-2 alleles in nature are nonrandomly associated with alleles at closely linked loci. If linkage disequilibrium is present in the experiments because it exists in nature, then the observed effects (such as frequency-dependent selection) would affect the Mdh-2 locus in nature as well.     

Drought and cold spells trigger dieback of temperate oak and beech forests in northern Spain

Dieback in temperate forests is understudied, despite this biome is predicted to be increasingly affected by more extreme climate events in a warmer world. To evaluate the potential drivers of dieback we reconstructed changes in radial growth and intrinsic water-use efficiency (iWUE) from stable isotopes in tree rings. Particularly, we compared tree size, radial-growth trends, growth responses to climate (temperature, precipitation, cloudiness, number of foggy days) and drought, and changes in iWUE of declining and non-declining trees showing contrasting canopy dieback and defoliation. This comparison was done in six temperate forests located in northern Spain and based on three broadleaved tree species (Quercus robur, Quercus humilis, Fagus sylvatica). Declining trees presented lower radial-growth rates than their non-declining counterparts and tended to show lower growth variability, but not in all sites. The growth divergence between declining and non-declining trees was significant and lasted more in Q. robur (15–30 years) than in F. sylvatica (5–10 years) sites. Dieback was linked to summer drought and associated atmospheric patterns, but in the wettest Q. robur sites cold spells contributed to the growth decline. In contrast, F. sylvatica was the species most responsive to summer drought in terms of growth reduction followed by Q. humilis which showed coupled changes in growth and iWUE as a function of tree vigour. Low growth rates and higher iWUE characterized declining Q. robur and F. sylvatica trees. However, declining F. sylvatica trees became less water-use efficient close to the dieback onset, which could indicate impending tree death. In temperate forests, dieback and growth decline can be triggered by climate extremes such as dry and cold spells, and amplified by climate warming and rising drought stress.     

Reproductive fitness,population size and genetic variation in Muscari tenuiflorum (Hyacinthaceae): The role of temporal variation

In plant populations a positive correlation between population size, genetic variation and fitness components is often found, due to increased pollen limitation or reduced genetic variation and inbreeding depression in smaller populations. However, components of fitness also depend on environmental factors which can vary strongly between years. The dry grassland species Muscari tenuiflorum experiences long term habitat isolation and small population sizes. We analyzed seed production of M. tenuiflorum in four years and its dependence on population size and genetic variation. Genetic diversity within populations was high (AFLP: H_e = 0.245; allozymes: H_e = 0.348). An analysis of molecular variance revealed considerable population differentiation (AFLP: 26%; allozyme: 17%). An overall pattern of isolation by distance was found, which, however was not present at distances below 20 km, indicating stronger effects of genetic drift. Genetic diversity was positively correlated to population size. Self pollination reduced seed set by 24%, indicating inbreeding depression. Reproductive fitness was not correlated to genetic diversity and a positive correlation with population size was present in two of four study years. The absence of a general pattern stresses the importance for multi-year studies. Overall, the results show that despite long term habitat isolation M. tenuiflorum maintains seed production in many years independent of population size. The long term persistence of populations is thus expected to depend less on intrinsic genetic or demographic properties affecting seed production but on successful plant establishment and persistence, which latter are based on conservation and protection of suitable habitats.     

Declining trend of Plasmodium falciparum dihydrofolate reductase (dhfr) and dihydropteroate synthase (dhps) mutant alleles after the withdrawal of Sulfadoxine-Pyrimethamine in North Western Ethiopia

Antimalarial drug resistance is one of the major challenges in global efforts of malaria control and elimination. In 1998, chloroquine was abandoned and replaced with sulfadoxine/pyrimethamine, which in turn was replaced with artemether/lumefantrine for the treatment of uncomplicated falciparum malaria in 2004. Sulfadoxine/pyrimethamine resistance is associated with mutations in dihydrofolate reductase (Pfdhfr) and dihydropteroate synthase (Pfdhps) genes. The prevalence of mutation in Pfdhfr and Pfdhps genes were evaluated and compared for a total of 159 isolates collected in two different time points, 2005 and 2007/08, from Pawe hospital, in North Western Ethiopia. The frequency of triple Pfdhfr mutation decreased significantly from 50.8% (32/63) to 15.9% (10/63) (P<0.001), while Pfdhps double mutation remained high and changed only marginally from 69.2% (45/65) to 55.4% (40/65) (P = 0.08). The combined Pfdhfr/Pfdhps quintuple mutation, which is strongly associated with sulfadoxine/pyrimethamine resistance, was significantly decreased from 40.7% (24/59) to 13.6% (8/59) (P<0.0001). On the whole, significant decline in mutant alleles and re-emergence of wild type alleles were observed. The change in the frequency is explained by the reduction of residual drug-resistant parasites caused by the strong drug pressure imposed when sulfadoxine/pyrimethamine was the first-line drug, followed by lower fitness of these resistant parasites in the absence of drug pressure. Despite the decrease in the frequency of mutant alleles, higher percentages of mutation remain prevalent in the study area in 2007/08 in both Pfdhfr and Pfdhps genes. Therefore, further multi-centered studies in different parts of the country will be required to assess the re-emergence of sulfadoxine/pyrimethamine sensitive parasites and to monitor and prevent the establishment of multi drug resistant parasites in this region.     

Balanced polymorphisms of quasineutral alleles

When the average number of progeny born to the genotypes at a locus are linear functions of a random variable such that, on the average, the number of progeny produced by each genotype is the same (quasineutrality), then there will be a systematic pressure on gene frequency that tends to favor the genotype with the smallest variance in the number of progeny. In particular, there will be a stable equilibrium attained whenever the variance in the number of progeny produced by the heterozygous genotype is less than that of both homozygous genotypes. We assume that the random variable affecting fitness is some function of an underlying environmental variable that tends to smooth out the day-to-day fluctuations in the environmental variable, such as the mean or the range or the variance. The effect of the environment on fitness can, therefore, be assumed to be approximately constant for periods corresponding to one generation. If the logarithm of the average number of progeny is a linear function of a random variable, then in contrast to the foregoing conclusions, there will be no tendency for any change in gene frequency at the locus (Kimura, M. (1954), Process leading to quasi-fixation of genes in natural populations due to random fluctuations of selection intensities, Genetics39, 280–295; (1964), Diffusion models in population genetics, J. App. Prob.1, 177–232.).     

Integrating demography and distribution modeling for the iconic Leontopodium alpinum Colm. in the Romanian Carpathians

1. Both climate change and human exploitation are major threats to plant life in mountain environments. One species that may be particularly sensitive to both of these stressors is the iconic alpine flower edelweiss (Leontopodium alpinum Colm.). Its populations have declined across Europe due to over‐collection for its highly prized flowers. Edelweiss is still subject to harvesting across the Romanian Carpathians, but no study has measured to what extent populations are vulnerable to anthropogenic change.
2. Here, we estimated the effects of climate and human disturbance on the fitness of edelweiss. We combined demographic measurements with predictions of future range distribution under climate change to assess the viability of populations across Romania.
3. We found that per capita and per‐area seed number and seed mass were similarly promoted by both favorable environmental conditions, represented by rugged landscapes with relatively cold winters and wet summers, and reduced exposure to harvesting, represented by the distance of plants from hiking trails. Modeling these responses under future climate scenarios suggested a slight increase in per‐area fitness. However, we found plant ranges contracted by between 14% and 35% by 2050, with plants pushed into high elevation sites.
4. Synthesis. Both total seed number and seed mass are expected to decline across Romania despite individual edelweiss fitness benefiting from a warmer and wetter climate. More generally, our approach of coupling species distribution models with demographic measurements may better inform conservation strategies of ways to protect alpine life in a changing world.

     

Spectral and statistical analysis of fluctuations in the SOL and diverted plasmas of the Uragan-3M torsatron

In the l = 3/m = 9 Uragan-3M (U3-M) torsatron (R ₀ = 1 m, ā ≈ 0.12 m, ι(ā)/2π ～ 0.3) with an open helical divertor and a plasma produced and heated by RF fields (ω ? ω_ci), studies of low frequency (5–100 kHz) density and potential fluctuations in the SOL plasma and in the diverted plasma flows (DPFs), have been carried out. It is shown, that in the SOL to more (less) distantly located points relative to the last closed magnetic surface, higher (lower) frequency fluctuations are inherent. Such a spectral splitting in two sub-ranges occurs in the DPFs too. A peculiarity of the spatial distribution of DPF fluctuation spectra is that lower frequency fluctuations dominate on the ion toroidal B × ?B drift side. During L-H-like transition in U-3M simultaneously with strong E _r shear formation, a suppression of lower frequency fluctuations and a decrease of local radial turbulent particle flux take place. Results are presented of investigation of plasma density fluctuations in SOL with the use of probability distribution function (PDF) analysis. Evaluations of skewness and kurtosis of fluctuations have been made. The analysis of I _s fluctuations in DPF have been carried out in a similar way.     

Effect of virulence genes complementary to the effective resistance genes in sorghum on fitness of the greenbug, <Emphasis Type="Italic">Schizaphis graminum</Emphasis> Rondani (Homoptera,Aphididae)

Analysis of the greenbug (Schizaphis graminum Rond.) — sorghum interaction system confirmed the hypothesis that rare insect virulence was related to reduced fitness. Greenbug clones from the Krasnodar population virulent to resistance genes Sgr5 and Sgr6 revealed lower fecundity in comparison with avirulent ones and were replaced in model populations during reproduction on a susceptible sorghum line. The main role of aphid fecundity was shown to provide higher fitness, reducing the frequency of virulent clones in natural populations.     

Kinship,parental manipulation and evolutionary origins of eusociality

One of the hallmarks of eusociality is that workers forego their own reproduction to assist their mother in raising siblings. This seemingly altruistic behaviour may benefit workers if gains in indirect fitness from rearing siblings outweigh the loss of direct fitness. If worker presence is advantageous to mothers, however, eusociality may evolve without net benefits to workers. Indirect fitness benefits are often cited as evidence for the importance of inclusive fitness in eusociality, but have rarely been measured in natural populations. We compared inclusive fitness of alternative social strategies in the tropical sweat bee, Megalopta genalis, for which eusociality is optional. Our results show that workers have significantly lower inclusive fitness than females that found their own nests. In mathematical simulations based on M. genalis field data, eusociality cannot evolve with reduced intra-nest relatedness. The simulated distribution of alternative social strategies matched observed distributions of M. genalis social strategies when helping behaviour was simulated as the result of maternal manipulation, but not as worker altruism. Thus, eusociality in M. genalis is best explained through kin selection, but the underlying mechanism is likely maternal manipulation.     

Negative frequency‐dependent selection maintains shell banding polymorphisms in two marine snails (Littorina fabalis and Littorina saxatilis)

The presence of shell bands is common in gastropods. Both the marine snails Littorina fabalis and Lttorina saxatilis are polymorphic for this trait. Such polymorphism would be expected to be lost by the action of genetic drift or directional selection, but it appears to be widespread at relatively constant frequencies. This suggests it is maintained by balancing selection on the trait or on a genetically linked trait. Using long time series of empirical data, we compared potential effects of genetic drift and negative frequency‐dependent selection (NFDS) in the two species. The contribution of genetic drift to changes in the frequency of bands in L. fabalis was estimated using the effective population size estimated from microsatellite data, while the effect of genetic drift in L. saxatilis was derived from previously published study. Frequency‐dependent selection was assessed by comparing the cross‐product estimator of fitness with the frequency of the polymorphism across years using a regression analysis. Both studied species showed patterns of NFDS. In addition, in L. fabalis, contributions from genetic drift could explain some of the changes in banding frequency. Overdominance and heterogeneous selection did not fit well to our data. The possible biological explanations resulting in the maintenance of the banding polymorphism are discussed.     

Freezing in a warming climate: Marked declines of a subnivean hibernator after a snow drought

Recent snow droughts associated with unusually warm winters are predicted to increase in frequency and affect species dependent upon snowpack for winter survival. Changes in populations of some cold‐adapted species have been attributed to heat stress or indirect effects on habitat from unusually warm summers, but little is known about the importance of winter weather to population dynamics and how responses to snow drought vary among sympatric species. We evaluated changes in abundance of hoary marmots (Marmota caligata) over a period that included a year of record‐low snowpack to identify mechanisms associated with weather and snowpack. To consider interspecies comparisons, our analysis used the same a priori model set as a concurrent study that evaluated responses of American pikas (Ochotona princeps) to weather and snowpack in the same study area of North Cascades National Park, Washington, USA. We hypothesized that marmot abundance reflected mechanisms related to heat stress, cold stress, cold exposure without an insulating snowpack, snowpack duration, atmospheric moisture, growing‐season precipitation, or select combinations of these mechanisms. Changes in marmot abundances included a 74% decline from 2007 to 2016 and were best explained by an interaction of chronic dryness with exposure to acute cold without snowpack in winter. Physiological stress during hibernation from exposure to cold, dry air appeared to be the most likely mechanism of change in marmot abundance. Alternative mechanisms associated with changes to winter weather, including early emergence from hibernation or altered vegetation dynamics, had less support. A post hoc assessment of vegetative phenology and productivity did not support vegetation dynamics as a primary driver of marmot abundance across years. Although marmot and pika abundances were explained by strikingly similar models over periods of many years, details of the mechanisms involved likely differ between species because pika abundances increased in areas where marmots declined. Such differences may lead to diverging geographic distributions of these species as global change continues.