On the relationship between food, reproduction and survival of two carabid beetles: Calathus melanocephalus and Pterostichus versicolor

Abstract.

• 1 The relative influences of temperature and availability of food on reproduction, survival and growth of all developmental stages of two carabid beetle species are discussed with special reference to the suggested relationship between availability of food, size of egg production and survival of adults from one breeding season to the next.
• 2 Temperature as well as food supply influence the length of larval growth and adult body size. Beetles grown at low temperatures and low amounts of food are smaller than those grown at higher temperature and with more food.
• 3 The number of eggs laid per female was correlated with the amount of food gathered. There was no inverse relationship (trade-off) between reproductive output and survival in the field until the next breeding season.
• 4 In 1980 no significant relationship was found between winter mortality and the amounts of food gathered by beetles in the period after reproduction and before winter diapause. However, in 1981 in C. melanocephalus a lower number of starved beetles survived the winter than the fed ones and‘field’beetles.
• 5 Only in the first part of the feeding activity period in autumn can enough food be gathered by C.melunocephalus for successful hibernation. In the second part of this period there is not enough food to build up the fat reserves needed to survive the winter.
• 6 Difference in population fluctuations of both species are discussed in relation to their life histories.

     

Patterns of movement of radioactive carabid beetles

Summary Tracking of individual ¹⁹²Ir-labeled ground beetles released in the field revealed that both the day-active and night-active species studied showed periods of small distances covered per day in random directions, alternating with periods of directed movement with large distances covered per day. This pattern occurred not only in the reproductive period but outside the breeding season as well in juvenile Pterostichus versicolor and spent Calathus melanocephalus. Although mean locomotory activity increased with temperature, great daily differences occurred between individuals, pointing to asynchronous behavior. In an unfavorable habitat directed movement occurred both more frequently and more extremely, sometimes resulting in escape to more favorable areas. Most of the radioactive beetles died within 7 weeks due to radiation effects, but independent field experiments and simulations showed that the recorded patterns were valid. Simulated individuals of P. versicolor living on 1 ha spread over 49 ha, whereas simulated C. melanocephalus covered only 9 ha after one activity season. Normal locomotory activities lead to both exchange of individuals between subpopulations and dispersal out of the habitat. The significance of these phenomena for population stability and for the survival of the species is discussed.Communication no. 184 of the Biological Station of the Agricultural University (Wageningen), Wijster (Drenthe), The Netherlands     

The significance of the diversity in age composition of Calathus melanocephalus L. (Col., Carabidae) in space and time at Schiermonnikoog

Summary Calathus melanocephalus L. reproduces at Schiermonnikoog in July, August and September. From literature it is known, that in some areas this species reproduces in August, September and October or still later.The population at Schiermonnikoog is composed of individuals of an old generation (adult overwintered beetles) which reproduce especially in July and in the first part of August and of individuals of a new generation (overwintered in the larval stage and hatched in June or July) which reproduce especially in the second part of August and in September.Both the differences in space and time in this age composition and those in fecundity are described.It is discussed, to what extent the differences in time of reproduction of populations in different localities in the same geographical area and of populations in different geographical areas, may be explained by the possible differences in age composition. In this context, differences between larval overwintering and adult overwintering species are discussed.The fecundity of the old females has been shown to be smaller than that of young females; moreover, the fecundity of individuals of both the old and new generation is different in the populations in the three localities studied.There is also pointed to the relation between age composition and the fecundity, with regard to the length of the reproductive period.Finally, the significance of the differences in space and time in age composition and fecundity to the chance of survival of Calathus melanocephalus in different areas is discussed.Communication of the Biological Station, Wijster, no. 160.     

Long adult life spain and polyphagy of a carabid beetle,Leptocarabus kumagaii in relation to reproduction and survival

The reproductive patterns of two different age classes of Leptocarabus kumagaii adults, i.e. young beetles which were in the first year of adult stage and old ones which survived for more than two years, were studied in the field. The numbers of eggs laid were compared between young and old females and the effects of food quality and quantity were studied on egg production and maintenance in the laboratory.

1. Both of the young and old beetles reproduced in the field population. The number of eggs laid did not significantly differ between the two.
2. The number of eggs laid by female increased with an increase in the amount of miced beef eaten. Plant materials were ineffective for egg production but sufficient for the survival of both male and female beetles.
3. The two life historical aspects, i.e. repeated reproduction and polyphagy were discussed in relation to environmental fluctuation and limitation of food resources.

     

Fecundity in relation to wing-morph of three closely related species of the melanocephalus group of the genus Calathus (Coleoptera: Carabidae)

Summary In two successive years the fecundity of the carabid beetles Calathus (Neocalathus) cinctus, C. (N.) melanocephalus and C. (N.) mollis was studied in relation to wing-morph and temperature. Differences were found between the three species in both egg production and timing and length of the oviposition period. In all species the fecundity of laboratory bred beetles was significantly higher than that of females collected in the field. Long-winged females of both cinctus and melanocephalus had significantly higher egg production than short-winged females, and they also tended to produce eggs over a longer period. In mollis only the fecundity of the long-winged morph was established. The observed lower relative fitness of the short-winged morph in both cinctus and melanocephalus contradicts the supposed increase of the frequency of this morph in ageing, more or less isolated, populations of these species. The loss of long-winged genotypes, resulting from flight activities, is considered the most plausible cause of the increase of short-winged beetles in ageing populations. The higher fecundity of macropterous females makes them especially suited for (re)establishing populations.Communication No. 429 of the Biological Station WijsterPresent address and address for offprint requests: Kortenburg 31, NL-6871 ND Rentum     

Demographic correlates of variable egg survival for a tropical lizard

Summary Anolis limifrons is a small iguanid lizard that is found in lowland rainforest in Central America. Important life attributes include early maturity (3–4 mo), multiple clutches of a single egg per breeding season, and low survival. Population turnover is essentially annual as less than 5% adults live more than one year. My objectives were to resolve the following specific questions: 1) Where are eggs laid? 2) Does survival differ among different types of oviposition site? 3) How is the survival of eggs related to the density and age distribution of lizards at different areas? Most eggs are laid beneath litter on the ground. Eggs laid aboveground are placed in accumulations of soil and litter in stumps, crevices in buttressed trees and vines, holes in trees, and on the tops of logs. Eggs were found in aboveground sites in proportion to the abundance of those sites. The major cause of mortality was predation by Solenopsis ants. Judging by the ratio of eggs known to have hatched to those that were predated, survival did not differ among the various types of oviposition site. Study areas with high egg survival in the middle of the wet season had relatively more young (<4–6 mo old) lizards at the end of the wet season than areas with low egg survival. The strong correlation between the relative number of young individuals and egg survival is a consequence of the seasonal cycle of growth and reproduction. Most eggs are laid in the wet season (May–December). Hatchlings complete growth during the dry season (January–April) and become reproductive adults in their second wet season. However, because hatchlings can grow to reproductive size in approximately 3 months, individuals that hatch early in the wet season may themselves produce offspring that will be <4–6 mo old in the late wet season. The relative survival of juveniles (eggs) and adults is an important datum for many theories of life history evolution. For A. limifrons, survival of eggs was two to three times more variable than survival of lizards on both temporal and spatial scales. Comparative data for reptiles are not available to put this information into an evolutionary prespective.     

The early‐life environment and individual plasticity in life‐history traits

We tested whether the early‐life environment can influence the extent of individual plasticity in a life‐history trait. We asked: can the early‐life environment explain why, in response to the same adult environmental cue, some individuals invest more than others in current reproduction? Moreover, can it additionally explain why investment in current reproduction trades off against survival in some individuals, but is positively correlated with survival in others? We addressed these questions using the burying beetle, which breeds on small carcasses and sometimes carries phoretic mites. These mites breed alongside the beetle, on the same resource, and are a key component of the beetle's early‐life environment. We exposed female beetles to mites twice during their lives: during their development as larvae and again as adults during their first reproductive event. We measured investment in current reproduction by quantifying average larval mass and recorded the female's life span after breeding to quantify survival. We found no effect of either developing or breeding alongside mites on female reproductive investment, nor on her life span, nor did developing alongside mites influence her size. In post hoc analyses, where we considered the effect of mite number (rather than their mere presence/absence) during the female's adult breeding event, we found that females invested more in current reproduction when exposed to greater mite densities during reproduction, but only if they had been exposed to mites during development as well. Otherwise, they invested less in larvae at greater mite densities. Furthermore, females that had developed with mites exhibited a trade‐off between investment in current reproduction and future survival, whereas these traits were positively correlated in females that had developed without mites. The early‐life environment thus generates individual variation in life‐history plasticity. We discuss whether this is because mites influence the resources available to developing young or serve as important environmental cues.     

The biology of the ground beetle Harpalus rufipes in a strawberry field in Northumberland

Harpalus rufipes (Degeer) was studied in a strawberry plot in Northumberland from 1973 to 1978 by pitfall trapping, and in the laboratory. Adults were active from April until November. Overwintered male beetles predominated at the beginning of each season until May, followed by overwintered females in June and July. Newly emerged, mainly female, beetles were active from August onwards. Overwintered females matured during early summer and laid eggs in August with a fecundity of 10–15 eggs/female. In the laboratory about 30% of beetles survived from one breeding season to the next. First- and second-instar larvae were caught in pitfall traps in autumn; in the laboratory they made approximately vertical burrows in which they stored seeds taken from the soil surface. Third-instar larvae fed on these seeds and were not active on the surface. Preferred seeds were those of grasses and Chenopodium album L. Larvae were usually aggregated in the soil at densities of 3–20/m².     

Pattern of movement in and dispersal from a dutch forest of Carabus problematicus Hbst. (Coleoptera,Carabidae)

Summary Data in the literature that in the Netherlands Carabus problematicus is an autumn breeder with overwintering larvae and adults have been confirmed. Pitfall catches show two peaks of adult beetles. The first small peak of activity is realized in May, the significance of which is not yet understood. The second higher peak occurs from the end of August till the beginning of October: the reproduction period. In this period at least three age classes are present. Some females were shown to take part in reproduction for at least two seasons. Old females appeared to have less eggs in the ovaries than young ones. Larvae can be trapped throughout the winter. The new generation appears in July and in the beginning of August. They immediately take part in reproduction. C. problematicus shows noctural activity with a peak towards the end of the night.In field experiments the pattern of movement and the dispersal behaviour of marked individuals was investigated and related to the choice of habitat. It was shown that in the Netherlands C. problematicus prefers a forest habitat. Some individuals were found to disperse from their favoured habitat. The percentage of beetles that disperse out of the forest during the breeding season could be estimated at 4–8%. Dispersal increases in the second part of the breeding season. Females tend to disperse more frequently than males.The pattern of movement consists of two types: a random walk and a directed movement. It is suggested that the pattern of movement of individuals dispersing out of the forest is the directed movement. It could be shown that individuals released outside the forest were able to walk towards a forest silhouette. The significance of both the pattern of movement and the dispersal behaviour for the survival of the species is discuussed.Communication no. 212 of the Biological Station of the Agricultural University (Wageningen), Wijster (Drente), The Netherlands     

All-year reproduction and possible thermal melanism in <Emphasis Type="Italic">Amblystogenium pacificum</Emphasis> (Coleoptera: Carabidae) on the sub-Antarctic Ile de la Possession (Iles Crozet)

In the fell-field habitat at 140–270 m altitude on the sub-Antarctic Ile de la Possession, Iles Crozet, the dimorphic beetle Amblystogenium pacificum Putzeys (Coleoptera: Carabidae) was studied monthly throughout the southern year 1993–1994 by timed hand collections involving turning stones on the rocky fell-field. There were many mating couples and females containing well-developed eggs in every month and the species reproduced throughout the year, with no evidence of a winter cessation of egg production. However, the frequency of mating couples was about twice as great in the summer half-year (October–March) as in the winter half (April–September). Relatively small differences in monthly mean temperatures between the summer and winter at Iles Crozet render all-year reproduction not unexpected when comparisons are made with the temperatures and phenology of Carabidae in cool habitats in other parts of the world. Mating pairs were collected at unusually high frequencies for carabids. The frequency of such pairs, taken per unit effort by hand collecting, was highest in October–December, lower from January to May and intermediate from June to September. Comparison of beetles taken as solitary, non-mating individuals with those taken as mating couples showed that in summer, females of the black morph of this dimorphic species were more frequently involved in mating than expected, but no differences were detected in winter. Female beetles were divided into younger individuals (putative age 1–2 years) and older females (putative age 3–5 years or more) by the amount of claw-wear on individuals of this species. This division indicated that about 40% of the older females showed no egg development when captured, but only about 20% of the younger females were non-breeding. However, of the reproducing beetles, young and old females matured similar numbers of eggs. An altitude transect showed that the proportion of the black morph increased significantly with altitude. Studies at 140–270 m showed that a significantly smaller proportion of the older females of the brown morph were breeding than in the younger brown individuals, and this effect was particularly evident in the colder winter period. No such age differences existed amongst the black morph.     

Timing of current reproduction directly affects future reproductive output in European coots

Life-history theory suggests that the variation in the seasonal timing of reproduction within populations may be explained on the basis of individual optimization. Optimal breeding times would vary between individuals as a result of trade-offs between fitness components. The existence of such trade-offs has seldom been tested empirically. We experimentally investigated the consequences of altered timing of current reproduction for future reproductive output in the European coot (Fulica atra). First clutches of different laying date were cross-fostered between nests, and parents thereby experienced a delay or an advance in the hatching date. The probability and success of a second brood, adult survival until and reproduction in the next season were then compared to the natural variation among control pairs. Among control pairs the probability of a second brood declined with the progress of season. Delayed pairs were less likely and advanced pairs were more likely to produce a second brood. These changes were quantitatively as predicted from the natural seasonal decline. The number of eggs in the second clutch was positively related to egg number in the first clutch and negatively related to laying date. Compared to the natural variation, delayed females had more and advanced females had fewer eggs in their second clutch. The size of the second brood declined with season, but there was no significant effect of delay or advance. Local adult survival was higher following a delay and reduced following an advance. The effect of the experiment on adult survival was independent of sex. Laying date and clutch size of females breeding in the next year were not affected by treatment. The study demonstrates the existence of a trade-off between increased probability of a second brood and decreased parental survival for early breeding. Timing-dependent effects of current reproduction on future reproductive output may thus play an important role in the evolution of the seasonal timing of reproduction.     

Magellanic penguin telomeres do not shorten with age with increased reproductive effort,investment, and basal corticosterone

All species should invest in systems that enhance longevity; however, a fundamental adult life‐history trade‐off exists between the metabolic resources allocated to maintenance and those allocated to reproduction. Long‐lived species will invest more in reproduction than in somatic maintenance as they age. We investigated this trade‐off by analyzing correlations among telomere length, reproductive effort and output, and basal corticosterone in Magellanic penguins (Spheniscus magellanicus). Telomeres shorten with age in most species studied to date, and may affect adult survival. High basal corticosterone is indicative of stressful conditions. Corticosterone, and stress, has been linked to telomere shortening in other species. Magellanic penguins are a particularly good model organism for this question as they are an unusually long‐lived species, exceeding their mass‐adjusted predicted lifespan by 26%. Contrary to our hypothesis, we found adults aged 5 years to over 24 years of age had similar telomere lengths. Telomeres of adults did not shorten over a 3‐year period, regardless of the age of the individual. Neither telomere length, nor the rate at which the telomeres changed over these 3 years, correlated with breeding frequency or investment. Older females also produced larger volume clutches until approximately 15 years old and larger eggs produced heavier fledglings. Furthermore, reproductive success (chicks fledged/eggs laid) is maintained as females aged. Basal corticosterone, however, was not correlated with telomere length in adults and suggests that low basal corticosterone may play a role in the telomere maintenance we observed. Basal corticosterone also declined during the breeding season and was positively correlated with the age of adult penguins. This higher basal corticosterone in older individuals, and consistent reproductive success, supports the prediction that Magellanic penguins invest more in reproduction as they age. Our results demonstrate that telomere maintenance may be a component of longevity even with increased reproductive effort, investment, and basal corticosterone.     

Cascading costs of reproduction in female house wrens induced to lay larger clutches

In many species, females produce fewer offspring than they are capable of rearing, possibly because increases in current reproductive effort come at the expense of a female's own survival and future reproduction. To test this, we induced female house wrens (Troglodytes aedon) to lay more eggs than they normally would and assessed the potential costs of increasing cumulative investment in the three main components of the avian breeding cycle – egg laying, incubation and nestling provisioning. Females with increased clutch sizes reared more offspring in the first brood than controls, but fledged a lower proportion of nestlings. Moreover, nestlings of experimental females were lighter than those of control females as brood size and prefledging mass were negatively correlated. In second broods of the season, when females were not manipulated, experimental females laid the same number of eggs as controls, but experienced an intraseasonal cost through reduced hatchling survival and a lower number of young fledged. Offspring of control and experimental females were equally likely to recruit to the breeding population, although control females produced more recruits per egg laid. The reproductive success of recruits from broods of experimental and control females did not differ. The manipulation also induced interseasonal costs to future reproduction, as experimental females had lower fecundity than controls when breeding at least 2 years after having their reproductive effort experimentally increased. Finally, females producing the modal clutch size of seven eggs in their first broods had the highest lifetime number of fledglings.     

Changes in life-cycle strategies of Carabus problematicus over a range of altitudes in Northern England

ABSTRACT.

• 1 Mandible tip length has been used to estimate the age of individual C.problematicus Herbst caught over an altitude range of more than 700 m in northern England.
• 2 At 100 m the great majority of females laid eggs in the summer of their emergence as adults. The female life-cycle is predominantly annual and 24% only of the egg-bearing individuals were in their second year.
• 3 Above 250 m, most females did not reproduce until the summer of the year following emergence, giving a biennial life-cycle. 96% of egg-bearing females caught above 830 m were in their second year.
• 4 All the males caught during the breeding season at 100 m had emerged within that calendar year and it is concluded that the male life-cycle is annual at this altitude.
• 5 Above 250 m, 83% of the males caught during the breeding season were in their first calendar year, suggesting that recently emerged males may inseminate second year females and also that the male life-cycle is predominantly annual over the whole altitude range studied.
• 6 At the higher temperatures of the lowland site females not only produce eggs in the year of emergence, they also produce more eggs per individual than at the higher sites. Egg production depends on food intake which involves mandible wear and individuals at the lowland site have significantly higher rate of mandible wear than at the sites above 250 m.

     

Early‐breeding females experience greater telomere loss

Annual reproductive success is often highest in individuals that initiate breeding early, yet relatively few individuals start breeding during this apparently optimal time. This suggests that individuals, particularly females who ultimately dictate when offspring are born, incur costs by initiating reproduction early in the season. We hypothesized that increases in the ageing rate of somatic cells may be one such cost. Telomeres, the repetitive DNA sequences on the ends of chromosomes, may be good proxies of biological wear and tear as they shorten with age and in response to stress. Using historical data from a long‐term study population of dark‐eyed juncos (Junco hyemalis), we found that telomere loss between years was greater in earlier breeding females, regardless of chronological age. There was no relationship between telomere loss and the annual number of eggs laid or chicks that reached independence. However, telomere loss was greater when temperatures were cooler, and cooler temperatures generally occur early in the season. This suggests that environmental conditions could be the primary cause of accelerated telomere loss in early breeders.     

A Model of Mutual Tolerance and the Origin of Communal Associations Between Unrelated Females

We use a genetic algorithm model employing game theory to explore the ecological conditions favoring reproductive tolerance between two unrelated females that meet at a nesting site (i.e., breeding resource). Each female adopts one of three strategies: (1) fight for exclusive use of the nest, (2) tolerate the other female and breed communally, but fight back if attacked, or (3) leave in search of new breeding opportunities. Nests vary in the number of offspring they can support and their probability of failure due to discovery by competitors. The model predicts that communal associations are most likely to arise when (1) the benefits of nest sharing to females exceed the losses to individual reproduction, (2) additional nesting sites are rare, (3) females have limited clutch sizes, and (4) dominant females are able to skew reproduction in their favor. The amount of reproductive skew a dominant (larger) female can acquire while maintaining a communal association is predicted to increase when the asymmetry in fighting ability between females increases, and at nests that have a relatively high probability of nesting success for solitary females. When the losers of fights can parasitize the winner's brood, dominant females must reduce reproductive skew to promote a communal relationship. We discuss the ability of our model to predict patterns of facultative communal behavior in burying beetles (Silphidae; Nicrophorus spp.), as well as the absence of communal behavior in dung beetles (Scarabaeidae).     

Benefits of extra food to reproduction depend on maternal condition

The amount of food resources available to upper‐level consumers can show marked variations in time and space, potentially resulting in food limitation. The availability of food resources during reproduction is a key factor modulating variation in reproductive success and life‐history tradeoffs, including patterns of resource allocation to reproduction versus self‐maintenance, ultimately impacting on population dynamics. Food provisioning experiments constitute a popular approach to assess the importance of food limitation for vertebrate reproduction. In this study of a mesopredatory avian species, the lesser kestrel Falco naumanni, we provided extra food to breeding individuals from egg laying to early nestling rearing. Extra food did not significantly affect adult body condition or oxidative status. However, it increased the allocation of resources to flight feathers moult and induced females to lay heavier eggs. Concomitantly, it alleviated the costs of laying heavier eggs for females in poor body condition, and reduced their chances of nest desertion (implying complete reproductive failure). Extra food provisioning improved early nestling growth (body mass and feather development). Moreover, extra food significantly reduced the negative effects of ectoparasites on nestling body mass, while fostering forearm (a flight apparatus trait) growth among highly parasitized nestlings. Our results indicate that lesser kestrels invested the extra food mainly to improve current reproduction, suggesting that population growth in this species can be limited by food availability during the breeding season. In addition, extra food provisioning reduced the costs of the moult–breeding overlap and affected early growth tradeoffs by mitigating detrimental ectoparasite effects on growth and enhancing development of the flight apparatus with high levels of parasitism. Importantly, our findings suggest that maternal condition is a major trait modulating the benefits of extra food to reproduction, whereby such benefits mostly accrue to low‐quality females with poor body condition.     

Modulation of the adrenocortical response to acute stress with respect to brood value, reproductive success and survival in the Eurasian hoopoe

Reproducing parents face the difficult challenge of trading-off investment in current reproduction against presumed future survival and reproduction. Glucocorticoids are supposed to mediate this trade-off because the adrenocortical response to stress disrupts normal reproductive behaviour in favour of self-maintenance and own survival. According to the brood-value hypothesis, individuals with a low survival probability until the next reproductive season have to invest in current reproduction, a process driven by a down-regulation of their adrenocortical response. If the adrenocortical response to stress effectively mediates the trade-off between current reproduction versus future survival and reproduction, we expect a negative relationship with reproductive success and a positive correlation of the adrenocortical stress response with survival. We studied the relationship between corticosterone secretion in parents and their current brood value, reproductive success and survival in a short-lived multi-brooded bird, the Eurasian hoopoe Upupa epops. The adrenocortical response to acute handling stress was correlated with the brood value within the individual (first and second broods of the year) and between individuals. Birds breeding late in the season mounted a lower total corticosterone response to acute stress than birds breeding earlier, while females showed lower levels than males. We observed a negative relationship between the adrenocortical stress response and rearing success or fledging success in females, as predicted by the brood-value hypothesis. However, we could not evidence a clear link between the adrenocortical stress response and survival. Future research testing the brood-value hypothesis and trade-offs between current reproduction and future survival should also measure free corticosterone and carefully differentiate between cross-sectional (i.e. between-individual) and individual-based experimental studies.     

Some aspects of the reproductive biology of Synodontis schall (Bloch-Schneider, 1801) from the White Nile near Khartoum

The biology of reproduction was investigated in 631 specimens of Synodontis schall taken from the White Nile near Khartoum. Morphological changes of the gonads were followed for twelve months and the results used to classify their different developmental stages and maturity. Water and ash content of the gonads varied slightly from month to month and this variation was more pronounced during the breeding season. Gonosomatic indices also varied in the same manner. Synodontis schall spawns from July to September and females are ripe at the early age of I. Absolute fecundity ranged from 10 000 to 90 000 eggs and was highly correlated with growth parameters such as length, weight and age. Variations in the diameter of the eggs during the breeding season were also followed.     

THE EFFECT OF BIRTH DATE ON FITNESS OF FEMALE DWARF PERCH,MICROMETRUS MINIMUS (PERCIFORMES: EMBIOTOCIDAE)

Lifetime reproductive success may vary considerably with birth date. I measured phenotypic selection on female birth date in a viviparous teleost fish (Embiotocidae: Micrometrus minimus) by sampling birth-date cohorts over time in Tomales Bay, California. Four episodes of selection were measured: survival from birth to first reproduction, reproductive success in the first breeding season, survival to second reproduction, and reproductive success in the second season. Birth date had a significant impact on fitness in the first two episodes. Early born females were more successful in their first breeding season than late born females (directional selection on birth date), but early born females were less likely to survive the period between birth and first reproduction, relative to females born in the middle of the season (stabilizing selection on birth date). The final two episodes of selection had no detectable effect on birth date. Because of the relationship between birth date and survival in the first year, overall selection on female birth date was stabilizing.