Effect of winter cold duration on spring phenology of the orange tip butterfly,Anthocharis cardamines

The effect of spring temperature on spring phenology is well understood in a wide range of taxa. However, studies on how winter conditions may affect spring phenology are underrepresented. Previous work on Anthocharis cardamines (orange tip butterfly) has shown population‐specific reaction norms of spring development in relation to spring temperature and a speeding up of post‐winter development with longer winter durations. In this experiment, we examined the effects of a greater and ecologically relevant range of winter durations on post‐winter pupal development of A. cardamines of two populations from the United Kingdom and two from Sweden. By analyzing pupal weight loss and metabolic rate, we were able to separate the overall post‐winter pupal development into diapause duration and post‐diapause development. We found differences in the duration of cold needed to break diapause among populations, with the southern UK population requiring a shorter duration than the other populations. We also found that the overall post‐winter pupal development time, following removal from winter cold, was negatively related to cold duration, through a combined effect of cold duration on diapause duration and on post‐diapause development time. Longer cold durations also lead to higher population synchrony in hatching. For current winter durations in the field, the A. cardamines population of southern UK could have a reduced development rate and lower synchrony in emergence because of short winters. With future climate change, this might become an issue also for other populations. Differences in winter conditions in the field among these four populations are large enough to have driven local adaptation of characteristics controlling spring phenology in response to winter duration. The observed phenology of these populations depends on a combination of winter and spring temperatures; thus, both must be taken into account for accurate predictions of phenology.     

Latitudinal variation in thermal reaction norms of post‐winter pupal development in two butterflies differing in phenological specialization

Latitudinal clines in thermal reaction norms of development are a common phenomenon in temperate insects. Populations from higher latitudes often develop faster throughout the range of relevant temperatures (i.e countergradient variation) because they must be able to complete their life cycle within a shorter seasonal time window compared to populations at lower latitudes. In the present study, we experimentally demonstrate that two species of butterflies Anthocharis cardamines (L.) and Pieris napi (L.) instead show a cogradient variation in thermal reaction norms of post‐winter pupal development so that lower latitude populations develop faster than higher latitude populations. The two species share host plants but differ in the degree of phenological specialization, as well as in the patterns of voltinism. We suggest that the pattern in A. cardamines, a univoltine phenological specialist feeding exclusively on flowers and seedpods, is the result of selection for matching to the phenological pattern of its local host plants. The other species, P. napi, is a phenological generalist feeding on the leaves of the hosts and it shows a latitudinal cline in voltinism. Because the latitudinal pattern in P. napi was an effect of slow development in a fraction of the pupae from the most northern population, we hypothesize that this population may include both bivoltine and univoltine genotypes. Consequently, although the two species both showed cogradient patterns in thermal reaction norms, it appears likely that this was for different reasons. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 981–991.     

Population abundance gradient of Inurois punctigera along altitude

A winter geometrid moth, Inurois punctigera, shows sympatric and genetically isolated seasonal populations (i.e. early‐ and late‐winter populations) in the cold regions of Japan, whereas it shows only mid‐winter populations in the warm regions. Variation in adult flight phenology on a large geographic scale along latitudinal environmental gradients has been described, but the phenological variation on a more local scale along altitudinal environmental gradients has not yet been characterized. In the present study, we assessed the flight phenology at high‐ and low‐elevation areas in Mt. Rokko, Hyogo, Japan. First, we revealed that flight period was not disrupted in mid‐winter, even at high‐elevation areas (>660 m) but the population abundance was much lower in high‐elevation areas than in low‐elevation areas. Then, in the following two seasons, we investigated I. punctigera abundance, winter harshness (i.e. winter temperature) and their host plant abundance in nine closely located stations in Mt. Rokko. A generalized linear mixed model analysis indicated a greater effect of winter temperature on I. punctigera abundance compared to available food resources, suggesting that differences in winter harshness among elevation shapes the gradient of I. punctigera abundance along altitude. Our findings suggest that harsh conditions during winter function as selective agents on mid‐winter types of I. punctigera, and this could be involved in the divergence between sympatric early‐ and late‐winter populations of I. punctigera.     

Genetic and environmental influences on leaf phenology and cold hardiness of native and introduced riparian trees

To explore the roles of plasticity and genetic variation in the response to spatial and temporal climate variation, we established a common garden consisting of paired collections of native and introduced riparian trees sampled along a latitudinal gradient. The garden in Fort Collins, Colorado (latitude 40.6°N), included 681 native plains cottonwood (Populus deltoides subsp. monilifera) and introduced saltcedar (Tamarix ramosissima, T. chinensis and hybrids) collected from 15 sites at 29.2–47.6°N in the central United States. In the common garden both species showed latitudinal variation in fall, but not spring, leaf phenology, suggesting that the latitudinal gradient in fall phenology observed in the field results at least in part from inherited variation in the critical photoperiod, while the latitudinal gradient in spring phenology observed in the field is largely a plastic response to the temperature gradient. Populations from higher latitudes exhibited earlier bud set and leaf senescence. Cold hardiness varied latitudinally in both fall and spring for both species. For cottonwood, cold hardiness began earlier and ended later in northern than in southern populations. For saltcedar northern populations were hardier throughout the cold season than southern populations. Although cottonwood was hardier than saltcedar in midwinter, the reverse was true in late fall and early spring. The latitudinal variation in fall phenology and cold hardiness of saltcedar appears to have developed as a result of multiple introductions of genetically distinct populations, hybridization and natural selection in the 150 years since introduction.     

Climate warming: a loss of variation in populations can accompany reproductive shifts

The most documented response of organisms to climate warming is a change in the average timing of seasonal activities (phenology). Although we know that these average changes can differ among species and populations, we do not know whether climate warming impacts within‐population variation in phenology. Using data from five study sites collected during a 13‐year survey, we found that the increase in spring temperatures is associated with a reproductive advance of 10 days in natural populations of common lizards (Zootoca vivipara). Interestingly, we show a correlated loss of variation in reproductive dates within populations. As illustrated by a model, this shortening of the reproductive period can have significant negative effects on population dynamics. Consequently, we encourage tests in other species to assess the generality of decreased variation in phenological responses to climate change.     

Genetic signs of multiple colonization events in Baltic ciscoes with radiation into sympatric spring‐ and autumn‐spawners confined to early postglacial arrival

Presence of sympatric populations may reflect local diversification or secondary contact of already distinct forms. The Baltic cisco (Coregonus albula) normally spawns in late autumn, but in a few lakes in Northern Europe sympatric autumn and spring‐ or winter‐spawners have been described. So far, the evolutionary relationships and taxonomic status of these main life history forms have remained largely unclear. With microsatellites and mtDNA sequences, we analyzed extant and extinct spring‐ and autumn‐spawners from a total of 23 Swedish localities, including sympatric populations. Published sequences from Baltic ciscoes in Germany and Finland, and Coregonus sardinella from North America were also included together with novel mtDNA sequences from Siberian C. sardinella. A clear genetic structure within Sweden was found that included two population assemblages markedly differentiated at microsatellites and apparently fixed for mtDNA haplotypes from two distinct clades. All sympatric Swedish populations belonged to the same assemblage, suggesting parallel evolution of spring‐spawning rather than secondary contact. The pattern observed further suggests that postglacial immigration to Northern Europe occurred from at least two different refugia. Previous results showing that mtDNA in Baltic cisco is paraphyletic with respect to North American C. sardinella were confirmed. However, the inclusion of Siberian C. sardinella revealed a more complicated pattern, as these novel haplotypes were found within one of the two main C. albula clades and were clearly distinct from those in North American C. sardinella. The evolutionary history of Northern Hemisphere ciscoes thus seems to be more complex than previously recognized.     

Latitudinal population differentiation in phenology,life history and flower morphology in the perennial herb Lythrum salicaria

Abstract In plants with a wide distribution, phenological characters can be expected to vary clinally along climatic gradients, whereas other characters important for adaptation to local biotic and abiotic factors may vary in a more mosaic fashion. We used common‐garden experiments and controlled crosses to examine population differentiation in phenology, life history and morphology in the perennial herb Lythrum salicaria along a latitudinal transect through Sweden (57°N to 66°N). Northern populations initiated growth and flowering earlier, flowered for a shorter period, were shorter, produced more and larger winter buds, and were older at first reproduction than southern populations. Flower morphology varied significantly among populations, but was, with the exception of calyx length, not significantly related to latitude of origin. Survival in the common garden (at 63°49′N) was positively correlated with latitude of origin and the size and number of winter buds produced in the preceding year. The results suggest that the among‐population differences in phenology and life history have evolved in response to latitudinal variation in length of the growing season. Further studies are required to determine whether population differentiation in flower morphology is maintained by selection.     

Seasonal distribution of the long-nosed bat (Leptonycteris curasoae) in North America: does a generalized migration pattern really exist?

Aim This paper examines the migration of the tropical nectarivorous bat Leptonycteris curasoae considered as a latitudinal migrant that breeds in south-west United States and northern Mexico in spring and migrates southward during fall. We tested the hypothesis that the latitudinal migration occurs only locally given by the local availability of bat resources, leading to migratory movements in zones with seasonal scarcity of resources and to resident bat populations where resources are available throughout the year. Localization We analysed the presence of L. curasoae along its distribution range in North America (between 14°N and 33°N). Study cases were also conducted in three Mexican localities: the Tehuacán Valley (17°48′–18°58′N and 96°48′–97°43′W), Sonoran Desert (28°41′N and 110°15′W), and the coast of Jalisco (19°32′N and 105°07′W). Methods Geographic evidence for latitudinal migration of L. curasoae was analysed using 94-year capture records housed in twenty-two collections of North America. Records were analysed using a Geographical Information System (GIS), in which floral resources and capture records were integrated. Monthly captures in the Tehuacán Valley were conducted during three years and bats abundance and reproductive status were correlated with the phenology of bat resources. Bat captures were also conducted during two consecutive years in an extratropical desert during winter and spring, and during one spring in the coast of Jalisco. Results The latitudinal migration of L. curasoae in North America only occurs at latitudes near 30°N, whereas bats may be residents at latitudes lower than 21°N. Captures were associated always to the availability of floral resources in both geographical and local scales. Main conclusions The existence of resident populations in the tropics with two reproductive events support the hypothesis that migration only occurs in the northern distribution limit of this nectar-feeding bat.     

Evolution,plasticity and evolving plasticity of phenology in the tree species Alnus glutinosa

Both traits and the plasticity of these traits are subject to evolutionary change and therefore affect the long‐term persistence of populations and their role in local communities. We subjected clones from 12 different populations of Alnus glutinosa, located along a latitudinal gradient, to two different temperature treatments, to disentangle the distribution of genetic variation in timing of bud burst and bud burst plasticity within and among genotypes, populations, and regions. We calculated heritability and evolvability estimates for bud burst and bud burst plasticity and assessed the influence of divergent selection relative to neutral drift. We observed higher levels of heritability and evolvability for bud burst than for its plasticity, whereas the total phenological heritability and evolvability (i.e. combining timing of bud burst and bud burst plasticity) suggest substantial evolutionary potential with respect to phenology. Earlier bud burst was observed for the low‐latitudinal populations than for the populations from higher latitudes, whereas the high‐latitudinal populations did not show the expected delayed bud burst. This countergradient variation can be due to evolution towards increased phenological plasticity at higher latitudes. However, because we found little evidence for adaptive differences in phenological plasticity across the latitudinal gradient, we suggest differential frost tolerance as the most likely explanation for the observed phenological patterns in A. glutinosa.     

Tracking climate impacts on the migratory monarch butterfly

Understanding the impacts of climate on migratory species is complicated by the fact that these species travel through several climates that may be changing in diverse ways throughout their complete migratory cycle. Most studies are not designed to tease out the direct and indirect effects of climate at various stages along the migration route. We assess the impacts of spring and summer climate conditions on breeding monarch butterflies, a species that completes its annual migration cycle over several generations. No single, broad‐scale climate metric can explain summer breeding phenology or the substantial year‐to‐year fluctuations observed in population abundances. As such, we built a Poisson regression model to help explain annual arrival times and abundances in the Midwestern United States. We incorporated the climate conditions experienced both during a spring migration/breeding phase in Texas as well as during subsequent arrival and breeding during the main recruitment period in Ohio. Using data from a state‐wide butterfly monitoring network in Ohio, our results suggest that climate acts in conflicting ways during the spring and summer seasons. High spring precipitation in Texas is associated with the largest annual population growth in Ohio and the earliest arrival to the summer breeding ground, as are intermediate spring temperatures in Texas. On the other hand, the timing of monarch arrivals to the summer breeding grounds is not affected by climate conditions within Ohio. Once in Ohio for summer breeding, precipitation has minimal impacts on overall abundances, whereas warmer summer temperatures are generally associated with the highest expected abundances, yet this effect is mitigated by the average seasonal temperature of each location in that the warmest sites receive no benefit of above average summer temperatures. Our results highlight the complex relationship between climate and performance for a migrating species and suggest that attempts to understand how monarchs will be affected by future climate conditions will be challenging.     

Amphibian breeding phenology trends under climate change: predicting the past to forecast the future

Global climate warming is predicted to hasten the onset of spring breeding by anuran amphibians in seasonal environments. Previous data had indicated that the breeding phenology of a population of Fowler's Toads (Anaxyrus fowleri) at their northern range limit had been progressively later in spring, contrary to generally observed trends in other species. Although these animals are known to respond to environmental temperature and the lunar cycle to commence breeding, the timing of breeding should also be influenced by the onset of overwintering animals’ prior upward movement through the soil column from beneath the frost line as winter becomes spring. I used recorded weather data to identify four factors of temperature, rainfall and snowfall in late winter and early spring that correlated with the toads’ eventual date of emergence aboveground. Estimated dates of spring emergence of the toads calculated using a predictive model based on these factors, as well as the illumination of the moon, were highly correlated with observed dates of emergence over 24 consecutive years. Using the model to estimate of past dates of spring breeding (i.e. retrodiction) indicated that even three decades of data were insufficient to discern any appreciable phenological trend in these toads. However, by employing weather data dating back to 1876, I detected a significant trend over 140 years towards earlier spring emergence by the toads by less than half a day/decade, while, over the same period of time, average annual air temperature and annual precipitation had both increased. Changes in the springtime breeding phenology for late‐breeding species, such as Fowler's Toads, therefore may conform to expectations of earlier breeding under global warming. Improved understanding of the environmental cues that bring organisms out of winter dormancy will enable better interpretation of long‐term phenological trends.     

Maintenance of clonal diversity in Dipsa bifurcata (Fallén, 1810) (Diptera: Lonchopteridae). I. Fluctuating seasonal selection moulds long-term coexistence

The deterministic maintenance of clonal diversity in thelytokous taxa can be seen as a model for understanding how environmental heterogeneity both can stabilize genetic diversity and can allow coexistence of competing species. We here analyze the temporal fluctuations in clonal diversity in the thelytokous Lonchopterid fly, Dipsa bifurcata (Fallén, 1810), at four localities in Sweden over an 8-year period. Estimated fitness values for clones are cyclical, synchronous among populations and correlated with seasonal changes in the environment. Differential winter viability and emergence from overwintering along with differential reproductive rate during the summer appear to be the selective mechanisms by which long-term clonal diversity is maintained. In a companion paper (Tomiuk et al, 2004), we present a model for the maintenance of clonal diversity through the mechanism of differential diapause among clones, utilizing fitness values estimated from the data presented here. In general, our results imply that fluctuating seasonal fitnesses can maintain stable genetic polymorphism within populations, as well as coexistence between closely related competitors, when coupled with differences in diapause phenology.     

Evolution of butterfly seasonal plasticity driven by climate change varies across life stages

Photoperiod is a common cue for seasonal plasticity and phenology, but climate change can create cue–environment mismatches for organisms that rely on it. Evolution could potentially correct these mismatches, but phenology often depends on multiple plastic decisions made during different life stages and seasons that may evolve separately. For example, Pararge aegeria (Speckled wood butterfly) has photoperiod-cued seasonal life history plasticity in two different life stages: larval development time and pupal diapause. We tested for climate change-associated evolution of this plasticity by replicating common garden experiments conducted on two Swedish populations 30 years ago. We found evidence for evolutionary change in the contemporary larval reaction norm—although these changes differed between populations—but no evidence for evolution of the pupal reaction norm. This variation in evolution across life stages demonstrates the need to consider how climate change affects the whole life cycle to understand its impacts on phenology.     

Predicting bird phenology from space: satellite‐derived vegetation green‐up signal uncovers spatial variation in phenological synchrony between birds and their environment

Population‐level studies of how tit species (Parus spp.) track the changing phenology of their caterpillar food source have provided a model system allowing inference into how populations can adjust to changing climates, but are often limited because they implicitly assume all individuals experience similar environments. Ecologists are increasingly using satellite‐derived data to quantify aspects of animals' environments, but so far studies examining phenology have generally done so at large spatial scales. Considering the scale at which individuals experience their environment is likely to be key if we are to understand the ecological and evolutionary processes acting on reproductive phenology within populations. Here, we use time series of satellite images, with a resolution of 240 m, to quantify spatial variation in vegetation green‐up for a 385‐ha mixed‐deciduous woodland. Using data spanning 13 years, we demonstrate that annual population‐level measures of the timing of peak abundance of winter moth larvae (Operophtera brumata) and the timing of egg laying in great tits (Parus major) and blue tits (Cyanistes caeruleus) is related to satellite‐derived spring vegetation phenology. We go on to show that timing of local vegetation green‐up significantly explained individual differences in tit reproductive phenology within the population, and that the degree of synchrony between bird and vegetation phenology showed marked spatial variation across the woodland. Areas of high oak tree (Quercus robur) and hazel (Corylus avellana) density showed the strongest match between remote‐sensed vegetation phenology and reproductive phenology in both species. Marked within‐population variation in the extent to which phenology of different trophic levels match suggests that more attention should be given to small‐scale processes when exploring the causes and consequences of phenological matching. We discuss how use of remotely sensed data to study within‐population variation could broaden the scale and scope of studies exploring phenological synchrony between organisms and their environment.     

Life-cycle regulation and life history plasticity in the speckled wood butterfly: are reaction norms predictable?

We investigated whether interpopulational variation in life-cycle regulation and life-history plasticity, in response to photoperiod, is predictable from considerations of what would be the adaptive life cycle and life history in a given environment. The investigation was performed on five populations of the speckled wood butterfly, Pararge aegeria (L.) (Lepidoptera: Nymphalidae), from central and south Sweden, England, Spain and Madeira. Insects from all five populations were reared at all daylengths from 10 h to 20 h at 17^oC. Larval and pupal development times were noted. Predictions were met regarding the type of life-cycle regulation and the shape of reaction norms. Evidence for diapause (larval summer and winter diapause, pupal winter diapause) was found in the three northern populations (P. a. tircis) but not in the two southern populations (P. a. aegeria). Photoperiodic thresholds for diapause induction followed the predicted latitudinal patterns, and this was also the case regarding quantitative regulation of development time (by photoperiod) among directly developing individuals. Under direct development, development time was progressively shorter in shorter daylengms in the two Swedish populations, where this signals progressively later dates. This was not found in the English, Spanish and Madeiran populations where such a response is likely to be maladaptive, because one or more generations of larvae are present before summer solstice. There were also unexpected results, for which we propose preliminary adaptive explanations.     

Evolution and disappearance of sympatric Coregonus albula in a changing environment—A case study of the only remaining population pair in Sweden

During the past 50 years, Fennoscandian populations of spring‐spawning Baltic cisco (Coregonus albula), sympatric to common autumn‐spawners, have declined or disappeared; for example, three out of four known spring‐spawning populations in Sweden are regarded as extinct. Over the same period, the climate has changed and populations have been subject to other anthropogenic stressors. We compared historic (1960s) and recent (1990–2000s) morphological data from the still‐existent sympatric cisco populations in Lake Fegen, Sweden. Phenotypic changes were found for spring‐spawners making them more similar to the sympatric autumn‐spawners that had remained virtually unchanged. Based on results for other salmoniform fishes, a phenotypically plastic response to increased temperature during early development appears unlikely. The recent material was also analyzed with microsatellite markers; long‐term effective population size in spring‐spawners was estimated to be about 20 times lower than autumn‐spawners, with signs of long‐term gene flow in both directions and a recent genetic bottleneck in spring‐spawners. We suggest the change toward a less distinct phenotype in spring‐spawners to reflect a recent increase in gene flow from autumn‐spawners. Time since divergence was estimated to only c. 1,900 years (95% CI: 400–5,900), but still the Fegen populations represent the most morphologically and genetically distinct sympatric populations studied. Consequently, we hypothesize that less distinct population pairs can be even younger and that spring‐spawning may have repeatedly evolved and disappeared in several lakes since the end of the last glaciation, concurrent with changed environmental conditions.     

Functional analysis of Normalized Difference Vegetation Index curves reveals overwinter mule deer survival is driven by both spring and autumn phenology

Large herbivore populations respond strongly to remotely sensed measures of primary productivity. Whereas most studies in seasonal environments have focused on the effects of spring plant phenology on juvenile survival, recent studies demonstrated that autumn nutrition also plays a crucial role. We tested for both direct and indirect (through body mass) effects of spring and autumn phenology on winter survival of 2315 mule deer fawns across a wide range of environmental conditions in Idaho, USA. We first performed a functional analysis that identified spring and autumn as the key periods for structuring the among-population and among-year variation of primary production (approximated from 1 km Advanced Very High Resolution Radiometer Normalized Difference Vegetation Index (NDVI)) along the growing season. A path analysis showed that early winter precipitation and direct and indirect effects of spring and autumn NDVI functional components accounted for 45% of observed variation in overwinter survival. The effect size of autumn phenology on body mass was about twice that of spring phenology, while direct effects of phenology on survival were similar between spring and autumn. We demonstrate that the effects of plant phenology vary across ecosystems, and that in semi-arid systems, autumn may be more important than spring for overwinter survival.     

Seasonality of North Atlantic phytoplankton from space: impact of environmental forcing on a changing phenology (1998–2012)

Seasonal pulses of phytoplankton drive seasonal cycles of carbon fixation and particle sedimentation, and might condition recruitment success in many exploited species. Taking advantage of long‐term series of remotely sensed chlorophyll a (1998–2012), we analyzed changes in phytoplankton seasonality in the North Atlantic Ocean. Phytoplankton phenology was analyzed based on a probabilistic characterization of bloom incidence. This approach allowed us to detect changes in the prevalence of different seasonal cycles and, at the same time, to estimate bloom timing and magnitude taking into account uncertainty in bloom detection. Deviations between different sensors stressed the importance of a prolonged overlap between successive missions to ensure a correct assessment of phenological changes, as well as the advantage of semi‐analytical chlorophyll algorithms over empirical ones to reduce biases. Earlier and more intense blooms were detected in the subpolar Atlantic, while advanced blooms of less magnitude were common in the Subtropical gyre. In the temperate North Atlantic, spring blooms advanced their timing and decreased in magnitude, whereas fall blooms delayed and increased their intensity. At the same time, the prevalence of locations with a single autumn/winter bloom or with a bimodal seasonal cycle increased, in consonance with a poleward expansion of subtropical conditions. Changes in bloom timing and magnitude presented a clear signature of environmental factors, especially wind forcing, although changes on incident photosynthetically active radiation and sea surface temperature were also important depending on latitude. Trends in bloom magnitude matched changes in mean chlorophyll a during the study period, suggesting that seasonal peaks drive long‐term trends in chlorophyll a concentration. Our results link changes in North Atlantic climate with recent trends in the phenology of phytoplankton, suggesting an intensification of these impacts in the near future.     

Rising temperature and development in dragonfly populations at different latitudes

1. For modelling the future ecological responses to climate change, data on individual species and on variation within and between populations from different latitudes are required. 2. We examined life cycle regulation and growth responses to temperature in Mediterranean and temperate populations of a widespread European odonate, Orthetrum cancellatum. In an experiment, offspring from individual females from different parts of the range were kept separately to elucidate differences between families. 3. The experiment was run outdoors at 52°N at a natural photoperiod for almost a year. We used four temperature regimes, ambient (i.e. following local air temperature) and ambient temperature increased by 2, 4 and 6 °C, to mimic future temperature rise. A mathematical model was used to categorise the type of seasonal regulation and estimate parameters of the temperature response curve. 4. Growth rate varied significantly with temperature sum, survival and geographic origin, as well as with family. Offspring of all females from the temperate part of the range had a life cycle with a 12 h day‐length threshold necessary to induce diapause (i.e. diapause was induced once day length fell below 12 h). By contrast, Mediterranean families had a 10 h threshold or had an unregulated life cycle allowing winter growth. The temperature response did not significantly differ between populations, but varied between families with a greater variation in the optimum temperature for growth in the Mediterranean population. 5. The variation in seasonal regulation leads to a diversity in voltinism patterns within species, ranging from bivoltine to semivoltine along a latitudinal gradient. Given that the type of seasonal regulation is genetically fixed, rising temperatures will not allow faster than univoltine development in temperate populations. We discuss the consequences of our results in the light of rising temperature in central Europe.     

CLINAL VARIATION IN SEED TRAITS INFLUENCING LIFE CYCLE TIMING IN ARABIDOPSIS THALIANA

Early life‐history transitions are crucial determinants of lifetime survival and fecundity. Adaptive evolution in early life‐history traits involves a complex interplay between the developing plant and its current and future environments. We examined the plant's earliest life‐history traits, dissecting an integrated suite of pregermination processes: primary dormancy, thermal induction of secondary dormancy, and seasonal germination response. We examined genetic variation in the three processes, genetic correlations among the processes, and the scaling of germination phenology with the source populations’ climates. A spring annual life history was associated with genetic propensities toward both strong primary dormancy and heat‐induced secondary dormancy, alone or in combination. Lineages with similar proportions of winter and spring annual life history have both weak primary dormancy and weak thermal dormancy induction. A genetic bias to adopt a spring annual strategy, mediated by rapid loss of primary dormancy and high thermal dormancy induction, is associated with a climatic gradient characterized by increasing temperature in summer and rainfall in winter. This study highlights the importance of considering combinations of multiple genetically based traits along a climatic gradient as adaptive strategies differentiating annual plant life‐history strategies. Despite the genetic‐climatic cline, there is polymorphism for life‐history strategies within populations, classically interpreted as bet hedging in an unpredictable world.