Variation In Reproductive Timing Research Articles

Genetic variation in reproductive timing in a long-lived herbaceous perennial.

Reproductive fitness of individual plants depends on the timing of flowering, especially in mate-limited populations, such as those in fragmented habitats. When flowering time traits are associated with differential reproductive success, the narrow-sense heritability (h2 ) of traits will determine how rapidly trait means evolve in response to selection. Heritability of flowering time is documented in many annual plants. However, estimating h2 of flowering time in perennials presents additional methodological challenges, often including paternity assignment and trait expression over multiple years. We evaluated the h2 of onset and duration of flowering using offspring-midparent regressions and restricted maximum likelihood methods in an experimental population of an iterocarpic, perennial, herbaceous plant, Echinacea angustifolia, growing in natural conditions. We assessed the flowering time of the parental cohort in 2005 and 2006; the offspring in 2014 through 2017. We also examined the effects of the paternity assignment from Cervus and MasterBayes on estimates of h2 . We found substantial h2 for onset and duration of flowering. We also observed variation in estimates among years. The most reliable estimates for both traits fell in the range of 0.1-0.17. We found evidence of a genotype by year interaction for onset of flowering and strong evidence that genotypes are consistent in their duration of flowering across years. Substantial heritabilities in this population imply the capacity for a response to natural selection, while also suggesting the potential for differential contributions to adaptive evolution among seasons.

Timing and synchrony of birth in Eurasian lynx across Europe.

The ecology and evolution of reproductive timing and synchrony have been a topic of great interest in evolutionary ecology for decades. Originally motivated by questions related to behavioral and reproductive adaptation to environmental conditions, the topic has acquired new relevance in the face of climate change. However, there has been relatively little research on reproductive phenology in mammalian carnivores. The Eurasian lynx (Lynx lynx) occurs across the Eurasian continent, covering three of the four main climate regions of the world. Thus, their distribution includes a large variation in climatic conditions, making it an ideal species to explore reproductive phenology. Here, we used data on multiple reproductive events from 169 lynx females across Europe. Mean birth date was May 28 (April 23 to July 1), but was ~10 days later in northern Europe than in central and southern Europe. Birth dates were relatively synchronized across Europe, but more so in the north than in the south. Timing of birth was delayed by colder May temperatures. Severe and cold weather may affect neonatal survival via hypothermia and avoiding inclement weather early in the season may select against early births, especially at northern latitudes. Overall, only about half of the kittens born survived until onset of winter but whether kittens were born relatively late or early did not affect kitten survival. Lynx are strict seasonal breeders but still show a degree of flexibility to adapt the timing of birth to surrounding environmental conditions. We argue that lynx give birth later when exposed to colder spring temperatures and have more synchronized births when the window of favorable conditions for raising kittens is shorter. This suggests that lynx are well adapted to different environmental conditions, from dry and warm climates to alpine, boreal, and arctic climates. This variation in reproductive timing will be favorable in times of climate change, as organisms with high plasticity are more likely to adjust to new environmental conditions.

Natural variation in reproductive timing and X-chromosome nondisjunction in Caenorhabditis elegans.

Caenorhabditis elegans hermaphrodites first produce a limited number of sperm cells, before their germline switches to oogenesis. Production of progeny then ensues until sperm is depleted. Male production in the self-progeny of hermaphrodites occurs following X-chromosome nondisjunction during gametogenesis, and in the reference strain increases with age of the hermaphrodite parent. To enhance our understanding of the reproductive timecourse in C. elegans, we measured and compared progeny production and male proportion during the early and late reproductive periods of hermaphrodites for 96 wild C. elegans strains. We found that the two traits exhibited natural phenotypic variation with few outliers and a similar reproductive timing pattern as previous reports. Progeny number and male proportion were not correlated in the wild strains, implying that wild strains with a large brood size did not produce males at a higher rate. We also identified loci and candidate genetic variants significantly associated with male-production rate in the late and total reproductive periods. Our results provide an insight into life-history traits in wild C. elegans strains.

Intra‐population variation in reproductive timing covaries with thermal plasticity of offspring performance in perch Percafluviatilis

Life history theory posits that organisms should time their reproduction to coincide with environmental conditions that maximize their fitness. Population-level comparisons have contributed important insights on the adaptive value of reproductive timing and its association to environmental variation. Yet, despite its central role to ecology and evolution, the causes and consequences of variation in reproductive timing among individuals within populations are poorly understood in vertebrates other than birds. Using a combination of observational field studies and a split-brood experiment, we investigated whether differences in breeding time were associated with changes in hatching success, reproductive allocation and reaction norms linking offspring performance to temperature within an anadromous Baltic Sea population of perch Perca fluviatilis. Field observations revealed substantial variation in reproductive timing, with the breeding period lasting almost 2months and occurring in temperatures ranging from 10 to 21℃. The hatching success of perch decreased as the reproductive season progressed. At the same time, the reproductive allocation strategy changed over the season, late breeders (the offspring of which were introduced into a high resource environment and increased predation pressure) produced more and smaller eggs that resulted in smaller larvae, compared with early breeders. The split-brood experiment in which eggs were incubated in different temperatures (10, 12, 15, 18°C) showed that differences in reproductive timing were associated with a change in the shape of the reaction norm linking offspring performance to water temperature indicative of adaptive phenotypic plasticity, with the offspring of early breeders performing best in low temperatures and the offspring of late breeders performing best in high temperatures. The seasonal changes in reproductive traits and the shape of the thermal performance suggest time-dependent adaptive differences among individuals within the population. Management actions aimed at preserving and restoring variation in the timing of reproductive events will thus likely also influence variation in associated life history traits and thermal performance curves, which could safeguard populations against environmental challenges and changes associated with exploitation and global warming.

Birth timing generates reproductive trade-offs in a non-seasonal breeding primate.

The evolutionary benefits of reproductive seasonality are often measured by a single-fitness component, namely offspring survival. Yet different fitness components may be maximized by different birth timings. This may generate fitness trade-offs that could be critical to understanding variation in reproductive timing across individuals, populations and species. Here, we use long-term demographic and behavioural data from wild chacma baboons (Papio ursinus) living in a seasonal environment to test the adaptive significance of seasonal variation in birth frequencies. We identify two distinct optimal birth timings in the annual cycle, located four-month apart, which maximize offspring survival or minimize maternal interbirth intervals (IBIs), by respectively matching the annual food peak with late or early weaning. Observed births are the most frequent between these optima, supporting an adaptive trade-off between current and future reproduction. Furthermore, infants born closer to the optimal timing favouring maternal IBIs (instead of offspring survival) throw more tantrums, a typical manifestation of mother-offspring conflict. Maternal trade-offs over birth timing, which extend into mother-offspring conflict after birth, may commonly occur in long-lived species where development from birth to independence spans multiple seasons. Our findings therefore open new avenues to understanding the evolution of breeding phenology in long-lived animals, including humans.

Exposure to artificial light at night accelerates but does not override latitude-dependent seasonal reproductive response in a North American songbird.

In the modern era of industrialization, illuminated nights have become a common defining feature of human-occupied environments, particularly cities. Artificial light at night (ALAN) imposes several known negative impacts on the neuroendocrine system, metabolism, and seasonal reproduction of species living in the wild. However, we know little about the impact of ALAN on populations of birds that either live year-round in the same location or move to different latitudes across seasons. To test whether ALAN has a differing impact on the reproductive timing of bird populations that winter in sympatry but breed at different latitudes, we monitored sedentary and migratory male dark-eyed juncos that were or were not exposed to low intensity (∼2.5±0.5 lux) ALAN. All groups were held in common conditions and day length was gradually increased to mimic natural day length changes (NDL). We assessed seasonal reproductive response from initiation to termination of the breeding cycle. As expected based on earlier research, the sedentary birds exhibited earlier gonadal recrudescence and terminated breeding later than the migratory birds. In addition, resident and migrant birds exposed to ALAN initiated gonadal recrudescence earlier and terminated reproductive events sooner as compared to their conspecifics experiencing NDL. Importantly, the difference in the reproductive timing of sedentary and migratory populations was maintained even when exposed to ALAN. This variation in the seasonal reproductive timing may likely have a genetic basis or be the result of early developmental effects imposed due to different light regimes related to the latitude of origin. This study reveals first that ALAN accelerated reproductive development across both migrants and residents and second that latitude-dependent variation in reproductive timing is maintained despite exposure to ALAN. These results corroborate a relationship between latitude, population, and reproductive timing while also revealing ALAN's impact on seasonal reproductive timing. This study reveals that, ALAN accelerated reproductive development but maintained latitude-dependent variation in reproductive timing across both migrant and resident bird populations.

Functional genetic diversity in an exploited marine species and its relevance to fisheries management.

The timing of reproduction influences key evolutionary and ecological processes in wild populations. Variation in reproductive timing may be an especially important evolutionary driver in the marine environment, where the high mobility of many species and few physical barriers to migration provide limited opportunities for spatial divergence to arise. Using genomic data collected from spawning aggregations of Pacific herring (Clupea pallasii) across 1600 km of coastline, we show that reproductive timing drives population structure in these pelagic fish. Within a specific spawning season, we observed isolation by distance, indicating that gene flow is also geographically limited over our study area. These results emphasize the importance of considering both seasonal and spatial variation in spawning when delineating management units for herring. On several chromosomes, we detected linkage disequilibrium extending over multiple Mb, suggesting the presence of chromosomal rearrangements. Spawning phenology was highly correlated with polymorphisms in several genes, in particular SYNE2, which influences the development of retinal photoreceptors in vertebrates. SYNE2 is probably within a chromosomal rearrangement in Pacific herring and is also associated with spawn timing in Atlantic herring (Clupea harengus). The observed genetic diversity probably underlies resource waves provided by spawning herring. Given the ecological, economic and cultural significance of herring, our results support that conserving intraspecific genetic diversity is important for maintaining current and future ecosystem processes.

Resource‐related variables drive individual variation in flowering phenology and mediate population‐level flowering responses to climate in an asynchronously reproducing palm

AbstractMany tropical plant species show wide intra‐population variation in reproductive timing, resulting in the protracted presence of flowering and fruiting individuals. Various eco‐evolutionary drivers have been proposed as ultimate causes for asynchronous phenology, yet little is known about the proximate factors that control reproductive onset among individuals or that influence the proportion of trees producing new inflorescences within a population. We employed a nine‐year phenological record from 178 individuals of the hyperdominant, asynchronously flowering canopy palm, Oenocarpus bataua (Arecaceae)¸ to assess whether resource‐related variables influence individual‐ and population‐level flowering phenology. Among individuals, access to sunlight increased rates of inflorescence production, while the presence of resource sinks related to current investment in reproduction—developing infructescences—reduced the probability of producing new inflorescences. At the population level, climate anomalies induced by El Niño Southern Oscillation (ENSO) affected the proportion of the population producing inflorescences through time. Moreover, the effects of ENSO anomalies on flowering patterns depended on the prevalence of developing infructescences in the population, with stronger effects in periods of low developing‐infructescence frequency. Taken together, these results suggest that resource‐related variables can drive phenological differences among individuals and mediate population‐level responses to larger‐scale variables, such as climate anomalies. Consequently, a greater focus on the role of resource levels as endogenous cues for reproduction might help explain the frequent aseasonal phenological patterns observed among tropical plants, particularly those showing high intra‐population asynchrony.Abstract in Spanish is available with online material.

Financial opportunity costs and deaths among close kin are independently associated with reproductive timing in a contemporary high-income society

Evolutionary demography predicts that variation in reproductive timing stems from socio-ecologically contingent trade-offs between current and future reproduction. In contemporary high-income societies, the costs and benefits of current reproduction are likely to vary by socioeconomic status (SES). Two influential hypotheses, focusing on the parenthood 'wage penalty', and responses to local mortality have separately been proposed to influence the timing of parenthood. Economic costs of reproduction (i.e. income loss) are hypothesized to delay fertility, especially among high childhood SES individuals who experience greater opportunities to build capital through advantageous education and career opportunities. On the other hand, relatively low childhood SES individuals experience higher mortality risk, which may favour earlier reproduction. Here, we examine both hypotheses with a representative register-based, multigenerational dataset from contemporary Finland (N = 47 678). Consistent with each hypothesis, the predicted financial cost of early parenthood was smaller, and mortality among close kin was higher for individuals with lower childhood SES. Within the same dataset, lower predicted adulthood income and more kin deaths were also independently associated with earlier parenthood. Our results provide a robust demonstration of how economic costs and mortality relate to reproductive timing. We discuss the implications of our findings for demographic theory and public policy.

Canalization of Seasonal Phenology in the Presence of Developmental Variation: Seed Dormancy Cycling in an Annual Weed.

Variation in the developmental timing in one life stage may ramify within and across generations to disrupt optimal phenology of other life stages. By focusing on a common mechanism of developmental arrest in plants-seed dormancy-we investigated how variation in flowering time influenced seed germination behavior and identified potential processes that can lead to canalized germination behavior despite variation in reproductive timing. We quantified effects of reproductive timing on dormancy cycling by experimentally manipulating the temperature during seed maturation and the seasonal timing of seed dispersal/burial, and by assessing temperature-dependent germination of un-earthed seeds over a seasonal cycle. We found that reproductive timing, via both seed-maturation temperature and the timing of dispersal, strongly influenced germination behavior in the weeks immediately following seed burial. However, buried seeds subsequently canalized their germination behavior, after losing primary dormancy and experiencing natural temperature and moisture conditions in the field. After the complete loss of primary dormancy, germination behavior was similar across seed-maturation and dispersal treatments, even when secondary dormancy was induced. Maternal effects themselves may contribute to the canalization of germination: first, by inducing stronger dormancy in autumn-matured seeds, and second by modifying the responses of those seeds to their ambient environment. Genotypes differed in dormancy cycling, with functional alleles of known dormancy genes necessary for the suppression of germination at warm temperatures in autumn through spring across multiple years. Loss of function of dormancy genes abolished almost all dormancy cycling. In summary, effects of reproductive phenology on dormancy cycling of buried seeds were apparent only as long as seeds retained primary dormancy, and a combination of genetically imposed seed dormancy, maternally induced seed dormancy, and secondary dormancy can mitigate variation in germination behavior imposed by variation in reproductive phenology.

Does light pollution alter daylength? A test using light loggers on free-ranging European blackbirds (Turdus merula).

Artificial light at night is one of the most apparent environmental changes accompanying anthropogenic habitat change. The global increase in light pollution poses new challenges to wild species, but we still have limited understanding of the temporal and spatial pattern of exposure to light at night. In particular, it has been suggested by several studies that animals exposed to light pollution, such as songbirds, perceive a longer daylength compared with conspecifics living in natural darker areas, but direct tests of such a hypothesis are still lacking. Here, we use a combination of light loggers deployed on individual European blackbirds, as well as automated radio-telemetry, to examine whether urban birds are exposed to a longer daylength than forest counterparts. We first used activity data from forest birds to determine the level of light intensity which defines the onset and offset of daily activity in rural areas. We then used this value as threshold to calculate the subjective perceived daylength of both forest and urban blackbirds. In March, when reproductive growth occurs, urban birds were exposed on average to a 49-min longer subjective perceived daylength than forest ones, which corresponds to a 19-day difference in photoperiod at this time of the year. In the field, urban blackbirds reached reproductive maturity 19 day earlier than rural birds, suggesting that light pollution could be responsible of most of the variation in reproductive timing found between urban and rural dwellers. We conclude that light at night is the most relevant change in ambient light affecting biological rhythms in avian urban-dwellers, most likely via a modification of the perceived photoperiod.

Variation in spawning time promotes genetic variability in population responses to environmental change in a marine fish.

The level of phenotypic plasticity displayed within a population (i.e. the slope of the reaction norm) reflects the short-term response of a population to environmental change, while variation in reaction norm slopes among populations reflects spatial variation in these responses. Thus far, studies of thermal reaction norm variation have focused on geographically driven adaptation among different latitudes, altitudes or habitats. Yet, thermal variability is a function of both space and time. For organisms that reproduce at different times of year, such variation has the potential to promote adaptive variability in thermal responses for critical early life stages. Using common-garden experiments, we examined the spatial scale of genetic variation in thermal plasticity for early life-history traits among five populations of endangered Atlantic cod (Gadus morhua) that spawn at different times of year. Patterns of plasticity for larval growth and survival suggest that population responses to climate change will differ substantially, with increasing water temperatures posing a considerably greater threat to autumn-spawning cod than to those that spawn in winter or spring. Adaptation to seasonal cooling or warming experienced during the larval stage is suggested as a possible cause. Furthermore, populations that experience relatively cold temperatures during early life might be more sensitive to changes in temperature. Substantial divergence in adaptive traits was evident at a smaller spatial scale than has previously been shown for a marine fish with no apparent physical barriers to gene flow (∼200 km). Our findings highlight the need to consider the impact of intraspecific variation in reproductive timing on thermal adaptation when forecasting the effects of climate change on animal populations.

Interannual variation in reproductive phenology in a riverine fish assemblage: implications for predicting the effects of climate change and altered flow regimes

Summary Freshwater fish commonly exhibit differences in reproductive seasonality within assemblages. However, the extent of variation in reproductive timing among years is less well understood, particularly in relation to environmental variation. Phenological shifts in response to environmental variation, including climate change, can differ among species and thereby change temporal resource partitioning (e.g. in the use of nursery habitat, dietary items, etc.) and potentially the strength of biotic interactions such as competition. To assess interannual variation in reproductive phenology and temporal resource partitioning in an arid‐land river, we collected a data set on young‐of‐year fish in four years within an overall span of 16 years, from 1995 to 2010. Fish species spawned in a consistent order across years; however, the onset of spawning was between 4 and 28 days earlier in 2008–2010 than in 1995. Spawning onset advanced more for later‐spawning species, resulting in reduced temporal partitioning across the assemblage. Based on these observations, we developed four alternate models of assemblage‐level responses to alteration of flow regime based on the magnitude of phenological shifts and corresponding temporal partitioning among species. Phenological data are consistent with only one of these models. Convergence and overlap in spawning time among species may decrease resource partitioning at a critical life stage and alter assemblage composition in recruitment‐driven systems like arid‐land rivers. Understanding factors driving convergence and overlap in spawning time among species is key to predicting effects of ongoing climate change and alteration of flow regimes on fish assemblages.

Short photoperiods attenuate central responses to an inflammogen

In most parts of the world, environmental conditions vary in a predictable seasonal manner. Thus, seasonal variation in reproductive timing and immune function has emerged in some species to cope with disparate seasonal demands. During the long days of spring and summer when food availability is high and thermoregulatory demands low, Siberian hamsters invest in reproduction, whereas during the harsh short days of winter hamsters divert energy away from reproductive activities and modify immune capabilities. Many seasonal adaptations can be recapitulated in a laboratory setting by adjusting day length (photoperiod). Early-life photoperiods are important sources of seasonal information and can establish an individual's developmental trajectory. Siberian hamsters housed under short days (SD; 8 h light/day) recover more rapidly than long-day (LD; 16 h light/day) hamsters from immune activation with lipopolysaccharide (LPS). SD hamsters attenuate fever response, reduce cytokine production, and abrogate behavioral responses following LPS injection. The mechanism by which SD Siberian hamsters attenuate febrile response remains unspecified. It is possible that periphery-to-brain communication of inflammatory signals is altered by exposure to photoperiod. Rather than testing photoperiod effects on each of the multiple routes by which immunological cues are communicated to the CNS, we administered LPS intracerebroventricularly (i.c.v.) following adolescent exposure to either 6 weeks of SD or LD. Injection of LPS i.c.v. led to a similar immune reaction in SD hamsters as previously reported with intraperitoneal injection. Short days attenuated the response to LPS with diminished fever spike and duration, as well as decreased locomotor inactivity. Furthermore, only LD hamsters demonstrated anhedonic-like behavior following LPS injection as evaluated by decreased preference for a milk solution. These results suggest that photoperiodic differences in response to infection are due in part to changes in central immune activation.

Pre-laying climatic cues can time reproduction to optimally match offspring hatching and ice conditions in an Arctic marine bird

Individuals breeding in seasonal environments are under strong selection to time reproduction to match offspring demand and the quality of the post-natal environment. Timing requires both the ability to accurately interpret the appropriate environmental cues, and the flexibility to respond to inter-annual variation in these cues. Determining which cues are linked to reproductive timing, what these cues are predicting and understanding the fitness consequences of variation in timing, is therefore of paramount interest to evolutionary and applied ecologists, especially in the face of global climate change. We investigated inter-annual relationships between climatic variation and the timing of reproduction in Canada's largest breeding population of Arctic common eiders (Somateria mollissima) in East Bay, Nunavut. Warmer spring temperatures predicted both earlier mean annual laying dates and the earlier ice-free conditions required by ducklings for post-natal growth. Warmer springs had higher variation in this temperature cue, and the population laying distribution became increasingly positively-skewed in warmer summers, potentially indicating that more low-quality females had the opportunity to commence laying in warmer years. Females that timed laying to match duckling hatching just prior to fully ice-free conditions obtained the highest duckling survival probability. Inter-annual data on repeated breeding attempts revealed that the individuals examined show a similar degree of laying flexibility in response to climatic variation; however, there was significant individual variation in the absolute timing of laying within an average year. This work sheds light on how reproductive timing is related to and influenced by variation in local climate and provides vital information on how climate-related variation in reproductive timing influence a fitness measure in an Arctic species. Results are especially relevant to future work in polar environments given that global climatic changes are predicted to be most intense at high latitudes.

Clock polymorphism in Pacific salmon: evidence for variable selection along a latitudinal gradient

Seasonal timing of life-history events is often under strong natural selection. The Clock gene is a central component of an endogenous circadian clock that senses changes in photoperiod (day length) and mediates seasonal behaviours. Among Pacific salmonids (Oncorhynchus spp.), seasonal timing of migration and breeding is influenced by photoperiod. To expand a study of 42 North American Chinook salmon (Oncorhynchus tshawytscha) populations, we tested whether duplicated Clock genes contribute to population differences in reproductive timing. Specifically, we examined geographical variation along a similar latitudinal gradient in the polyglutamine domain (PolyQ) of OtsClock1a and OtsClock1b among 53 populations of three species: chum (Oncorhynchus keta), coho (Oncorhynchus kisutch) and pink salmon (Oncorhynchus gorbuscha). We found evidence for variable selection on OtsClock1b that corresponds to latitudinal variation in reproductive timing among these species. We evaluated the contribution of day length and a freshwater migration index to OtsClock1b PolyQ domain variation using regression trees and found that day length at spawning explains much of the variation in OtsClock1b allele frequency among chum and Chinook, but not coho and pink salmon populations. Our findings suggest that OtsClock1b mediates seasonal adaptation and influences geographical variation in reproductive timing in some of these highly migratory species.

Microevolution of neuroendocrine mechanisms regulating reproductive timing in Peromyscus leucopus

A key question in the evolution of life history and in evolutionary physiology asks how reproductive and other life-history traits evolve. Genetic variation in reproductive control systems may exist in many elements of the complex inputs that can affect the hypothalamic-pituitary-gonadal (HPG) or reproductive axis. Such variation could include numbers and other traits of secretory cells, the amount and pattern of chemical message released, transport and clearance mechanisms, and the number and other traits of receptor cells. Selection lines created from a natural population of white-footed mice (Peromyscus leucopus) that contains substantial genetic variation in reproductive inhibition in response to short winter daylength (SD) have been used to examine neuroendocrine variation in reproductive timing. We hypothesized that natural genetic variation would be most likely to occur in the inputs to GnRH neurons and/or in GnRH neurons themselves, but not in elements of the photoperiodic pathway that would have pleiotropic effects on nonreproductive functions as well as on reproductive functions. Significant genetic variation has been found in the GnRH neuronal system. The number of GnRH neurons immunoreactive to an antibody to mature GnRH peptide under conditions maximizing detection of stained neurons was significantly heritable in an unselected control (C) line. Furthermore, a selection line that suppresses reproduction in SD (photoperiod responsive, R) had fewer IR-GnRH neurons than a selection line that maintains reproduction in SD (photoperiod nonresponsive, NR). This supports the hypothesis that genetic variation in characteristics of GnRH neurons themselves may be responsible for the observed phenotypic variation in reproduction in SD. The R and NR lines differ genetically in food intake and iodo-melatonin receptor binding, as well as in other characteristics. The latter findings are consistent with the hypothesis that genetic variation occurs in the nutritional and hormonal inputs to GnRH neurons. Genetic variation also exists in the phenotypic plasticity of responses to two combinations of treatments, (1) food and photoperiod, and (2) photoperiod and age, indicating genetic variation in individual norms of reaction within this population. Overall, the apparent multiple sources of genetic variation within this population suggest that there may be multiple alternative combinations of alleles for both the R and NR phenotypes. If that interpretation is correct, we suggest that this offers some support for the evolutionary "potential" hypothesis and is inconsistent with the evolutionary "constraint" and "symmorphosis" hypotheses for the evolution of complex neuroendocrine pathways.

Divergent timing of egg‐laying may maintain life history polymorphism in potentially multivoltine insects in seasonal environments

The length of the favourable season determines voltinism in insect populations. In some insects, there is variation in fecundity and timing of reproduction among females. If the length of the favourable season does not allow all offspring to develop into adults without diapause, the benefits of high early fecundity may outweigh the associated cost of low lifetime fecundity. We tested this by exploring mating frequencies of Pieris napi females along a latitudinal gradient in different generations. Pieris napi is a bivoltine butterfly, and genetically polyandrous females enjoy higher lifetime fecundity than monandrous ones. Polyandry is, however, coupled with a relatively low early fecundity. We found that monandrous females are more likely to produce an additional generation than polyandrous ones under conditions that allow production of only a partial summer generation. Monandrous females were also the first to emerge and slightly over-represented in the summer generation under conditions that allow the development of a complete summer generation. Further, a stochastic model shows that variation in the timing of reproduction between strategies is sufficient to explain the observed patterns. Thus, seasonality may counter-select against polyandry, or more generally against low early reproductive rate, and promote maintenance of polymorphism in life history strategies.

Impact of the invasive cane toad (Bufo marinus) on an Australian frog (Opisthodon ornatus) depends on minor variation in reproductive timing

Invasive species are widely viewed as unmitigated ecological catastrophes, but the reality is more complex. Theoretically, invasive species could have negligible or even positive effects if they sufficiently reduce the intensity of processes regulating native populations. Understanding such mechanisms is crucial to predicting ultimate ecological impacts. We used a mesocosm experiment to quantify the impact of eggs and larvae of the introduced cane toad (Bufo marinus) on fitness-related traits (number, size and time of emergence of metamorphs) of a native Australian frog species (Opisthodon ornatus). The results depended upon the timing of oviposition of the two taxa, and hence the life-history stages that came into contact. Growth and survival of O. ornatus tadpoles were enhanced when they preceded B. marinus tadpoles into ponds, and reduced when they followed B. marinus tadpoles into ponds, relative to when tadpoles of both species were added to ponds simultaneously. The dominant tadpole-tadpole interaction is competition, and the results are consistent with competitive priority effects. However, these priority effects were reduced or reversed when O. ornatus tadpoles encountered B. marinus eggs. Predation on toxic toad eggs reduced the survival of O. ornatus and B. marinus. The consequent reduction in tadpole densities allowed the remaining O. ornatus tadpoles to grow more rapidly and to metamorphose at larger body sizes (>60% disparity in mean mass). Thus, exposure to B. marinus eggs reduced the number of O. ornatus metamorphs, but increased their body sizes. If the increased size at metamorphosis more than compensates for the reduced survival, the effective reproductive output of native anurans may be increased rather than decreased by the invasive toad. Minor interspecific differences in the seasonal timing of oviposition thus have the potential to massively alter the impact of invasive cane toads on native anurans.

Seasonal gonadal recrudescence in song sparrows: Response to temperature cues

Day length predicts changing of seasons at mid-latitudes, but additional environmental cues (e.g., temperature, rainfall) give more precise information about timing of food peaks critical for offspring survival. We tested the effects of temperature on seasonal reproductive development of male song sparrows ( Melospiza melodia morphna) from two populations: the Western Washington coast (3 m) and Cascade Mountains (500–1220 m). Previous work has shown that the timing of gonadal recrudescence can differ between the two sites in the field by as much as two months. However, in the laboratory under identical controlled conditions, testes grow at the same rate. To test whether temperature alone could account for a portion of the variation we measured in the field, we captured birds from each site and held them in temperature controlled environmental chambers that mimicked temperatures experienced either in the mountains or on the coast. We increased day length on a natural schedule, and measured testis volume, levels of circulating androgens and prolactin, and song rates. Increasing day length stimulated gonadal growth in all groups. We found only modest effects of temperature on reproductive development. In the mountain birds colder, montane temperatures slowed rates of growth, delaying the onset of growth by one month. Since temperature changes more markedly during the early winter months in the mountains than on the coast, increasing temperature may be a more relevant cue in timing of reproduction in the mountain population. These data suggest that while temperature helps to explain some of the variation in reproductive timing of free living sparrows, another as yet untested cue in the field may play a more important role. Furthermore, our data suggest that individuals within the same species may rely on different proximate cues for reproductive timing depending on the specific habitat in which they live.