Seasonal Environments Research Articles

Shifts in risk sensitivity and resource availability alter fat stores for a large mammal following extreme winter conditions

Abstract For species that inhabit environments where resource availability may be unpredictable, balance of resource allocation to life‐history traits can have heightened consequences for survival, reproduction, and ultimately, fitness. Acquisition and allocation of energy to maintenance, capital gain and reproduction should be in tune with the landscape an animal inhabits—environmental severity, food availability and population size all influence the resources animals have and dictate the ways they should be allocated. In seasonal environments, animals that experience periods of extreme resource limitation (e.g. harsh winters) may favour allocation of resources to body reserves to secure their survival at the cost of reproduction (i.e. risk averse). In contrast, the same accumulation of body reserves may not be necessary to survive in relatively benign landscapes where instead, allocation to reproduction is favoured (i.e. risk prone). According to the theory of risk‐sensitive allocation of resources, when animals are exposed to unprecedented or life‐threatening conditions, they may shift resource allocation to favour building capital over allocation in reproduction to preempt against encountering another life‐threatening event in the future. Using data from a long‐term project on a highly site‐faithful and long‐lived species, mule deer (Odocoileus hemionus), we evaluated how a life‐threatening winter and the associated changes in resource availability resulting from a population reduction influenced how animals acquired and allocated energy to survival (i.e. fat accumulation). Per capita precipitation, and the associated reduction in population abundance after the severe winter, had a positive influence of accrual of fat over summer. After the extreme physiological stress of a hard winter, deer starting spring with low body reserves accumulated 2.8 percentage points more fat over summer compared with before the experience of a bad winter and had an increased probability of recruiting fewer offspring. Fat stores can interact with environment, life history and behaviour to influence survival during periods of resource scarcity. For a long‐lived herbivore, we documented shifts in risk tolerance associated with fat accrual in preparation for winter, supporting the notion that risk‐sensitive allocation of resources may be plastic—an essential adaptation for animals to cope with rapidly changing landscapes. Read the free Plain Language Summary for this article on the Journal blog.

The evolution of enclosed nesting in passerines is shaped by competition, energetic costs, and predation threat

Abstract Many avian species breed in enclosed nests that may provide better protection against predation and climatic conditions compared to open nests and are generally associated with larger clutch sizes and slower offspring growth. Here we show that different enclosed nesting strategies are each linked to behaviors with very different costs and benefits on a macroevolutionary scale. Using a detailed dataset of nest structure and location from the order Passeriformes, we employed phylogenetic comparative methods to evaluate (1) how predation, competition, design complexity, and energetic costs have shaped evolutionary transitions between different nesting strategies, and (2) whether these strategies also have distinct relationships with life-history traits. We find that flexible strategies (i.e., nesting in both open and enclosed sites) as well as energetically demanding strategies are evolutionarily unstable, indicating the presence of underlying ecological tradeoffs between antipredator protections, construction costs, and competition. We confirm that species with enclosed nests have larger clutch sizes and longer development and nestling periods compared to open nesters, but only species that construct enclosed nests rather than compete for preexisting cavities spend more time incubating and are concentrated in the tropics. Flexible strategies prevail in seasonal environments and are linked to larger clutches—but not longer development—compared to nesting in the open. Overall, our results suggest that predation, competition, and energetic costs affect the evolution of nesting strategies, but via distinct pathways, and that caution is warranted when generalizing about the functions of enclosed nest designs in birds.

Does early spring arrival lead to early nesting in a migratory shorebird? Insights from remote tracking

Advancing breeding phenology is a commonly observed response to climate warming among bird species, potentially in response to shifts in the phenology of key resources. However, for migratory birds breeding at high latitudes, their capacity to breed earlier may be constrained by the time available between arrival on the breeding grounds and nesting, especially for later‐arriving individuals. This may have consequences for productivity, as early laying is often linked to higher breeding success, particularly in such seasonal environments. We investigated how migratory arrival timing influences subsequent timing of breeding, in particular whether the time between arrival and laying (arrival–laying gap) varies with arrival date, and if later‐arriving individuals are able to catch up with early‐arriving conspecifics. We tracked individual Icelandic Black‐tailed Godwit Limosa limosa islandica with GPS and PTT tags for one to two complete breeding seasons between 2013 and 2022. After arrival in Iceland, most Godwits visited their breeding territory within 5 days, though this interval was longer for earlier‐arriving birds. The total gap between arrival and laying was also longer for earlier‐arriving birds, such that laying date did not vary significantly with arrival date. These results suggest that, despite individual consistency in migratory phenology, subsequent timing of nesting is probably influenced by other factors, such as mate arrival timing and/or annual variation in environmental conditions. Regular pre‐nesting visits to the breeding territory may indicate that Godwits are able to monitor such factors closely and to nest early when conditions allow, but a larger sample of individuals and years is needed to assess whether early‐arriving individuals contribute disproportionately to population‐level advances in breeding phenology. Widespread tracking with high temporal and spatial resolution helps improve our understanding of phenological variation during the breeding season and its consequences for productivity and variation in juvenile phenology.

Seasonal variation in glucose and insulin is modulated by food and temperature conditions in a hibernating primate.

Feast-fast cycles allow animals to live in seasonal environments by promoting fat storage when food is plentiful and lipolysis when food is scarce. Fat-storing hibernators have mastered this cycle over a circannual schedule, by undergoing extreme fattening to stockpile fuel for the ensuing hibernation season. Insulin is intrinsic to carbohydrate and lipid metabolism and is central to regulating feast-fast cycles in mammalian hibernators. Here, we examine glucose and insulin dynamics across the feast-fast cycle in fat-tailed dwarf lemurs, the only obligate hibernator among primates. Unlike cold-adapted hibernators, dwarf lemurs inhabit tropical forests in Madagascar and hibernate under various temperature conditions. Using the captive colony at the Duke Lemur Center, we determined fasting glucose and insulin, and glucose tolerance, in dwarf lemurs across seasons. During the lean season, we maintained dwarf lemurs under stable warm, stable cold, or fluctuating ambient temperatures that variably included food provisioning or deprivation. Overall, we find that dwarf lemurs can show signatures of reversible, lean-season insulin resistance. During the fattening season prior to hibernation, dwarf lemurs had low glucose, insulin, and HOMA-IR despite consuming high-sugar diets. In the active season after hibernation, glucose, insulin, HOMA-IR, and glucose tolerance all increased, highlighting the metabolic processes at play during periods of weight gain versus weight loss. During the lean season, glucose remained low, but insulin and HOMA-IR increased, particularly in animals kept under warm conditions with daily food. Moreover, these lemurs had the greatest glucose intolerance in our study and had average HOMA-IR values consistent with insulin resistance (5.49), while those without food under cold (1.95) or fluctuating (1.17) temperatures did not. Remarkably low insulin in dwarf lemurs under fluctuating temperatures raises new questions about lipid metabolism when animals can passively warm and cool rather than undergo sporadic arousals. Our results underscore that seasonal changes in insulin and glucose tolerance are likely hallmarks of hibernating mammals. Because dwarf lemurs can hibernate under a range of conditions in captivity, they are an emerging model for primate metabolic flexibility with implications for human health.

Spring temperature drives phenotypic selection on plasticity of flowering time.

In seasonal environments, a high responsiveness of development to increasing temperatures in spring can infer benefits in terms of a longer growing season, but also costs in terms of an increased risk of facing unfavourable weather conditions. Still, we know little about how climatic conditions influence the optimal plastic response. Using 22 years of field observations for the perennial forest herb Lathyrus vernus, we assessed phenotypic selection on among-individual variation in reaction norms of flowering time to spring temperature, and examined if among-year variation in selection on plasticity was associated with spring temperature conditions. We found significant among-individual variation in mean flowering time and flowering time plasticity, and that plants that flowered earlier also had a more plastic flowering time. Selection favoured individuals with an earlier mean flowering time and a lower thermal plasticity of flowering time. Less plastic individuals were more strongly favoured in colder springs, indicating that spring temperature influenced optimal flowering time plasticity. Our results show how selection on plasticity can be linked to climatic conditions, and illustrate how we can understand and predict evolutionary responses of organisms to changing environmental conditions.

Periodic Trajectories for HIV Dynamics in a Seasonal Environment With a General Incidence Rate

In this paper, we propose a five-dimensional nonlinear system of differential equations for the human immunodeficiency virus (HIV) including the B-cell functions with a general nonlinear incidence rate. The compartment of infected cells was subdivided into three classes representing the latently infected cells, the short-lived productively infected cells, and the long-lived productively infected cells. The basic reproduction number was established, and the local and global stability of the equilibria of the model were studied. A sensitivity analysis with respect to the model parameters was undertaken. Finally, some numerical simulations are presented to illustrate the theoretical findings.

Underground trees inhabit varied environmental extremes across the Afrotropics.

Geoxyles, a distinctive feature of Afrotropical savannas and grasslands, survive recurrent disturbances by resprouting subshrub branches from large below-ground woody structures. Underground trees are a type of geoxyle that independently evolved within woody genera of at least 40 plant families in Africa. The environmental limits and determinants of underground tree biogeography are poorly understood, with the relative influence of frost and fire debated in particular. We aim to quantify variability in the niche of underground tree species relative to their taller, woody tree/shrub congeners. Using occurrence records of four Afrotropical genera, Parinari (Chrysobalanaceae), Ozoroa (Anacardiaceae), Syzygium (Myrtaceae) and Lannea (Anacardiaceae), and environmental data of nine climate and disturbance variables, the biogeography and niche of underground trees are compared with their open and closed ecosystem congeners. Along multiple environmental gradients and in a multidimensional environmental space, underground trees inhabit significantly distinct and extreme environments relative to open and closed ecosystem congeners. Niche overlap is low among underground trees and their congeners, and also among underground trees of the four genera. Of the study taxa, Parinari underground trees inhabit hotter, drier and more seasonal environments where herbivory pressure is greatest. Ozoroa underground trees occupy relatively more fire-prone environments, while Syzygium underground trees sustain the highest frost frequency and occur in relatively wetter conditions with seasonal waterlogging. Lannea underground trees are associated with the lowest temperatures, highest precipitation, and varying exposure to disturbance. While underground trees exhibit repeated convergent evolution, varied environments shape the ecology and biogeography of this iconic plant functional group. The multiplicity of extreme environments related to fire, frost, herbivory and waterlogging that different underground tree taxa occupy, and the distinctiveness of these environments, should be recognized in the management of African grassy ecosystems.

Phenological dynamics of four populations of Handroanthus spongiosus in seasonally dry tropical forest in Brazil

The scarcity of phenological studies based on different populations of tropical forest trees limits seed management and collection for reforestation efforts. Precipitation is the primary factor driving tropical plant phenology in seasonal environments, although other environmental variables and plant traits may be associated. We examined the seasonality, synchrony, and intensities of the vegetative and reproductive phenophases of four populations of Handroanthus spongiosus, an endangered species, under similar climate regimes in a seasonally dry tropical forest, in northeastern Brazil. We expected to observe some divergence in the phenologies of the populations related to distinct functional traits selected for by differences in rainfall and soil properties. Mature trees (n = 87) were monitored during a three-year period. Seasonality was examined using circular statistics, and the influences of environmental variables on phenophases were investigated using generalized additive models. Variations in intensities and activity indices were identified among the different populations. Vegetative phenophases were seasonal, driven by precipitation and photoperiod, with leaf longevity of up to 7 months; budding peaked in February-March, while leaf fall peaked in April and October. The reproductive phenophases were found to be seasonal, during the rainy season (November to April), influenced by temperature and photoperiod. The slight divergences noted among the phenological behaviors of the populations were related to distinct functional traits (e.g., tree height, stem diameter) selected for by differences in certain environmental variables (rainfall volumes and soil properties). Given ongoing global climate changes, increases in leaf fall and reductions of flowering intensity, as verified here, will likely be observed.

Maladaptive plastic responses of flowering time to geothermal heating.

Phenotypic plasticity might increase fitness if the conditions under which it evolved remain unaltered, but becomes maladaptive if the environment no longer provides reliable cues for subsequent conditions. In seasonal environments, timing of reproduction can respond plastically to spring temperature, maximizing the benefits of a long season while minimizing the exposure to unfavorable cold temperatures. However, if the relationship between early spring temperatures and later conditions changes, the optimal response might change. In geothermally heated ecosystems, the plastic response of flowering time to springtime soil temperature that has evolved in unheated areas is likely to be non-optimal, because soil temperatures are higher and decoupled from air temperatures in heated areas. We therefore expect natural selection to favor a lower plasticity and a delayed flowering in these areas. Using observational data along a natural geothermal warming gradient, we tested the hypothesis that selection on flowering time depends on soil temperature and favors later flowering on warmer soils in the perennial Cerastium fontanum. In both study years, plants growing in warmer soils began flowering earlier than plants growing in colder soils, suggesting that first flowering date (FFD) responds plastically to soil temperature. In one of the two study years, selection favored earlier flowering in colder soils but later flowering in warmer soils, suggesting that the current level of plastic advance of FFD on warmer soils may be maladaptive in some years. Our results illustrate the advantages of using natural experiments, such as geothermal ecosystems, to examine selection in environments that recently have undergone major changes. Such knowledge is essential to understand and predict both ecological and evolutionary responses to climate warming.

Climatic factors affecting seasonal movements of giraffes (Giraffa camelopardalis) in a semi‐arid region of South Africa

AbstractTo the modern‐day conservationist, knowledge on giraffe (Giraffa camelopardalis) movement patterns is crucial for the effective management of this increasingly threatened species. The aim of this study was to determine the seasonal distances travelled by giraffes in a highly seasonal semi‐arid savanna area and to investigate how specific climatic factors influence their spatial distribution. Distributed over the 95,537‐ha reserve, we selected eight adult female giraffes that were equipped with radio transmitter collars using geographical positioning system devices to investigate their movement patterns over a 21‐month period in the Kalahari region of South Africa. Climatic data were collected using three weather stations at different locations in the study area. Giraffes moved significantly longer distances and for longer hours during the cool, dry season compared to other seasons. Both temperature and rainfall had statistically significant effects on distances travelled. The giraffes moved significantly longer distances with higher rainfall and temperature. They also moved significantly further during the middle of the day in all seasons, except winter. The reasons for these movement patterns are discussed with special emphasis on the influence of climatic factors on resource availability in a highly seasonal environment with periods of low resource availability.

Seasonal variation in trophic structure and community niche dynamics of an Arctic coastal community of marine vertebrates

Temporal variation in food web structure is widespread among highly seasonal environments, such as the Arctic, and is driven by changes in resource availability. Variation in resource availability can lead to species differences in diet composition, isotopic niche width, and trophic position (TP) across seasons. Here, we used tissue samples that represent two distinct turnover rates for diet (liver = shorter term, muscle = longer term) from 18 fish and three marine mammal species to investigate seasonal (i) variation in TPs within the Southampton Island marine ecosystem of Hudson Bay, (ii) variation in consumer isotopic niche width within this part of the food web, and (iii) variation in community niche dynamics among a fish and marine mammal community using stable carbon (δ13C) and nitrogen (δ15N) isotope analyses. Many zooplanktivores and piscivores increased in TP in summer (i.e., shorter-term turnover period), whereas benthic feeders dependent on ice-obligate prey decreased in TP. Most isotopic niche widths and community metrics (δ15N range, total ellipse area, mean centroid distance) were higher in liver than muscle. Our findings demonstrate seasonal changes in TPs, which suggests that Arctic communities may be differentially affected by longer ice-free periods and earlier onset of primary production due to accelerated climate change.

Local food availability and nonbreeding carry-over effects affect breeding propensity and success of a tundra-nesting predator, the Long-tailed Jaeger

Abstract Reproduction of long-distance migrants can be affected by local conditions on the breeding grounds as well as those encountered during the nonbreeding season through carry-over effects. We show that this is true in Long-tailed Jaegers (Stercorarius longicaudus) because individuals that spent less time flying at sea during winter had a high breeding propensity and a reduced pre-laying interval, but breeding propensity and nesting success were also positively associated with food abundance at the breeding site. This seabird switches from a marine to a terrestrial lifestyle to breed in summer in the Arctic, where it primarily feeds on lemmings. We monitored jaeger reproduction and lemming densities on Bylot Island in the Canadian Arctic for 16 years, and we used geolocator to study annual movements. We assessed whether movement parameters (travel distance, migration duration, phenology, and number of flying bouts inferred by saltwater immersions) during the nonbreeding season affected the breeding propensity, phenology, and success of individuals. We also examined whether cyclic lemming fluctuations influenced Long-tailed Jaeger reproduction and whether nesting success affected the phenology of their outbound migration. We found that increased time spent flying during winter and early arrival at the breeding site reduced breeding propensity. Moreover, spending less time flying during winter shortened the pre-laying period, and advancing laying date increased nesting success. Birds may thus face a trade-off to minimize the relative costs associated with arriving too early and breeding too late. Local food availability had a strong effect on reproduction because breeding propensity and nesting success increased sharply with lemming abundance. Failed breeders advanced their outbound migration by 10 days on average compared to successful ones, but migration duration was similar. Therefore, the unpredictability of the highly seasonal Arctic environment, especially fluctuating food abundance, appears to be a strong driver of reproduction that can modulate the strength of carry-over effects.

Maternal size in perch (Perca fluviatilis) influences the capacity of offspring to cope with different temperatures

Climate change causes earlier and warmer springs in seasonal environments and a higher incidence of extreme weather events. In aquatic environments, this changes the thermal conditions during spawning, and the thermal performance of eggs and embryos may determine the consequences of climate change on recruitment. In iteroparous species with indeterminate growth, the eggs produced by a given female in successive years will increase in size as the female grows larger and likely be exposed to different temperatures during incubation due to annual variation in spring phenology. Still, we know little about whether differences in maternal size impact the temperature-dependent performance and viability of the offspring. Here we utilised a thermal gradient laboratory experiment on Baltic Sea perch (Perca fluviatilis) to investigate how maternal size influence the temperature dependent hatching success of the offspring. The results uncovered a positive relationship between maternal size and average hatching success, but the shape of the relationship (reaction norm) linking hatching success to incubation temperature was independent of maternal size. However, we did find an association between maternal size and the variance (S.D. and CV) in hatching success across temperatures, with larger females producing offspring with maintained performance (less sensitive) across temperature treatments, indicative of flatter reaction norms and broader thermal niches. This suggests that maintaining the size distribution of fish populations, for instance through regulations of size-selective fisheries, may be important to aid the long-term productivity and viability of fish populations and ultimately conserve the function and services of ecosystems.

Xylem structure and hydraulic function in roots and stems of chaparral shrub species from high and low elevation in the Sierra Nevada, California.

Xylem structure and hydraulics were compared between individuals at lower and upper elevation distribution limits for five chaparral shrub species along a steep transect in the southern Sierra Nevada, California, USA. Higher-elevation plants experienced frequent winter freeze-thaw events and increased precipitation. We hypothesized that environmental differences would lead to xylem trait differences between high and low elevations, but predictions were complicated because both water stress (low elevation) and freeze-thaw events (high elevation) may select for similar traits, such as narrow vessel diameter. We found significant changes in the ratio of stem xylem area to leaf area (Huber value) between elevations, with more xylem area required to support leaves at low elevations. Co-occurring species significantly differed in their xylem traits, suggesting diverse strategies to cope with the highly seasonal environment of this Mediterranean-type climate region. Roots were more hydraulically efficient and more vulnerable to embolism relative to stems, potentially due to roots being buffered from freeze-thaw stress, which allows them to maintain wider diameter vessels. Knowledge of the structure and function of both roots and stems is likely important in understanding whole-plant response to environmental gradients.

Diverse strategies for tracking seasonal environmental niches at hemispheric scale

AbstractAimSpecies depend upon a constrained set of environmental conditions, or environmental niches, for survival and reproduction that are being increasingly perturbed or lost under rapid climatic change. Seasonal environments, which require species to withstand shifting conditions or track their niches via movement, can offer an important system to study the range of biological responses to potentially cope with climate change. Here, we develop a novel methodological framework to identify niche‐tracking strategies, including the tracking of niche position and breadth, using a uniquely well‐sampled system of 619 New World bird species.LocationWestern Hemisphere.Time period1980–2020.Major taxa studiedBirds (Aves).MethodsAt continental scales, we identify the tracking of both environmental niche position and breadth and assess its phylogenetic and functional underpinning. Partitioning niche position and breadth tracking can inform whether climatic means or extremes constrain seasonal niches.ResultsWe uncover four primary niche‐tracking strategies, including the tracking of environmental niche position, niche breadth, both or neither. Species that track niche position most often also track niche breadth, but nearly 40% only track one component and 26% only track niche breadth and not position. There is only limited phylogenetic determinism to this variation, but a strong association with ecological and functional attributes that differs between niche position versus niche breadth tracking.Main conclusionsThe observed diversity in type and strength of environmental niche‐tracking strategies points to highly differing sensitivity to ongoing climatic change, with narrow trackers of both position and breadth particularly susceptible. The trait associations of niche tracking imply significant functional consequences for communities and ecosystems as impending climate change affects some strategies more strongly than others. Seasonal environments and their diversity of niche‐tracking strategies offer exceptionally dynamic systems for understanding the biological responses and consequences of climate change.

Body size and its correlates in fruit-feeding butterflies in a seasonal environment

Body size and its correlates in fruit-feeding butterflies in a seasonal environment

Breeding of Lotus japonicus that can overcome adverse seasonal environment by coupling flowering time and abiotic stresses

AbstractManipulation of floral transition may allow for a free change in the duration of plant vegetative and reproductive growth, thus rendering them to avoid adverse seasonal environmental conditions. In this report, we investigated the effect of the stress‐inducible promoter RD29A on driving the rice FT ortholog Hd3a expression in Lotus japonicus ‘MG20’ in response to environmental changes. Constitutive overexpression of Hd3a (HOE) in ‘MG20’ hastened its flowering in contrast to non‐transformed control (wild type; WT), which did not flower in winter hampered by long days and low temperatures. RD29A promoter had low activity under non‐stress conditions and was induced by cold, drought and salinity in transgenic ‘MG20’. Transgenic plants of Hd3a driven by the RD29A promoter (RH) could flower under unfavourable weather conditions that prohibited the flowering of WT plants. In addition, RH transgenic plants exhibited thriving clumps of branching, inflorescence, and pods because of the low temperature‐induced expression of Hd3a. These results demonstrate that inducible expression of florigen has the potential to tackle unfavourable weather conditions and boost plant production.

The effects of seasonal changes on the dynamics of a fig tree's pollination

Fig trees and their pollinating wasps are mutually dependent on each other. Both partners' reproductive success is regulated by the capacity of fig wasps to enter receptive figs at an appropriate time for pollination and oviposition. Oviposition is dependent on successful female pollinator dispersal from one tree to another and although fig wasps are slow flyers and short-lived they can be carried long distances by the wind. The relative importance of local versus long-distance pollinator dispersal is unclear, as is how this may vary with season. In the highly seasonal environment of the Makana Botanical Gardens, Grahamstown, South Africa, we recorded fruiting phenologies of all the trees in a monoecious Ficus burtt-davyi Hutchinson population together with variation in the abundance of its pollinator Elisabethiella baijnathi Wiebes. By comparing captures of fig wasps flying in the air with the numbers that emerged locally, we also examined the independence of the fig tree population, which was separated from the nearest conspecifics by more than 1 km. The abundance of pollinators flying in the air and the number of fig wasps released by figs were correlated with temperature. During winter there were times when no pollinators were released locally; however, they were still caught in the traps, showing that the wasps had dispersed from elsewhere and that the population was not totally independent. These results highlight the ability of fig wasps to disperse between populations and the likely impact of seasonal fluctuations on fig tree gene flow.

Remote monitoring of short‐term body mass variation in savanna ungulates

AbstractLarge herbivores in seasonal environments often experience mass variation due to temporal changes in the availability of critical resources like water and forage, as well as due to breeding events. Yet the documentation of mass variation in mammals of highly seasonal savanna habitats, which host the highest densities of grazing ungulates globally, has rarely been explored. Here, we showcase a method to evaluate seasonal mass variation in bovids. Our method used mineral‐baited scales and camera traps to enable us to track the body mass of three species through a period of wet and dry seasons in a South African savanna ecosystem. To illustrate one potential application of the method, we related body mass data to time, weather and resource availability. This showed that individuals altered their body masses markedly between seasons with, for example, female Kudu (Tragelaphus strepsiceros) gaining, on average, >21 kg over the 15‐week wet‐season period in 1 year. These changes were positively related to factors such as vegetation productivity (assessed using NDVI) and the frequency of rains. This method enables easy, non‐lethal and non‐invasive acquisition of mass data. The equipment is easy to deploy concurrently over large areas. Monitoring by this method has a variety of possible applications, potentially providing a useful early‐warning indicator of body condition to inform management, or providing information about ecological states, such as parturition or the reproductive effort of males. Given the longer and harsher dry seasons experienced in many arid systems in recent decades, and projected in future, this method may provide a straightforward means of monitoring long‐term body condition in animals as a result of environmental change.

Seasonality and competition select for variable germination behavior in perennials.

The occurrence of within-population variation in germination behavior and associated traits such as seed size has long fascinated evolutionary ecologists. In annuals, unpredictable environments are known to select for bet-hedging strategies causing variation in dormancy duration and germination strategies. Variation in germination timing and associated traits is also commonly observed in perennials, and often tracks gradients of environmental predictability. Although bet-hedging is thought to occur less frequently in long-lived organisms, these observations suggest a role of bet-hedging strategies in perennials occupying unpredictable environments. We use complementary analytical and evolutionary simulation models of within-individual variation in germination behavior in seasonal environments to show how bet-hedging interacts with fluctuating selection, life-history traits, and competitive asymmetries among germination strategies. We reveal substantial scope for bet-hedging to produce variation in germination behavior in long-lived plants, when "false starts" to the growing season results in either competitive advantages or increased mortality risk for alternative germination strategies. Additionally, we find that lowering adult survival may, in contrast to classic bet-hedging theory, result in less spreading of germination by decreasing density-dependent competition. These models extend insights from bet-hedging theory to perennials and explore how competitive communities may be affected by ongoing changes in climate and seasonality patterns.