Clutch size and malarial parasites in female great tits

life-history models predict an evolutionary trade-off in theallocation of resources to current versus future reproduction.This corresponds, at the physiological level, to a trade-offin the allocation of resources to current reproduction or tothe immune system, which will enhance survival and thereforefuture reproduction. For clutch size, life-history models predicta positive correlation between current investment in eggs andthe subsequent parasite load. In a population of great tits,we analyzed the correlation between natural clutch size of femalesand the subsequent prevalence of Plasmodium spp., a potentiallyharmful blood parasite. Females that showed, 14 days after hatchingof the nestlings, an infection with Plasmodium had a significantlylarger clutch (9.3 eggs ± 0.5 SE, n = 18) than uninfectedfemales (8.0 eggs ± 0.2 SE, n = 80), as predicted bythe allocation trade-off. Clutch size was positively correlatedwith the prevalence of Plasmodium, but brood size 14 days afterhatching was not. This suggests that females incur higher costsduring laying the clutch than during rearing nestlings. Infectionstatus of some females changed between years, and these changeswere significantly correlated with a change in clutch size aspredicted by the trade-off. The link between reproductive effortand parasitism may represent a possible mechanism by which thecost of egg production is mediated into future survival andmay thereby be an important selective force in the shaping ofclutch size     

Reproductive effort reduces long-term immune function in breeding tree swallows (Tachycineta bicolor)

We examined whether strategies of reproductive allocation may reduce long-term immunocompetence through the effects of manipulated effort on secondary or acquired immunity. We tested whether increased reproductive effort leads to reduced immune function and survival by manipulating brood size in tree swallows (Tachycineta bicolor) and exposing breeding females to a primary and secondary exposure of sheep red blood cells to elicit a humoral immune response. Females raising enlarged broods produced fewer secondary antibodies than did females raising control or reduced broods. Most importantly, individuals with high secondary responses were more likely to survive to breed 3 years after brood manipulations, suggesting that differences in disease susceptibility may be caused by trade-offs in reproductive allocation. We also found that individual quality, measured by clutch initiation date, mediated the effects of brood manipulations, with higher-quality birds showing a greater ability to deal with increases in effort.     

Patterns of resource allocation in male and female plants of Rumex acetosa and R. acetosella

Summary Patterns of resource allocation in the dioecious Rumex acetosa and R. acetosella were investigated. Males were found to allocate more to reproduction during flower production than females, whereas females invested considerably more in reproduction during seed production. Altogether, females allocated both a higher total amount and a higher proportion of energy to reproduction than did males. By regression analysis, the influence of plant size on reproductive effort was examined separately for males and females. The results indicated that while reproductive effort is sometimes lower for tall plants than for small plants, size-independent effects have a greater influence on reproductive effort than size distribution. An analysis of variance was conducted to investigate the effects of population, season, sex and their interactions on plant size, and an analysis of covariance was used to study differences in resource allocation patterns. Different interaction effects were found to be most important in the two species of Rumex.     

A physiological marker for quantifying differential reproductive investment between the sexes in Yellow-legged gulls (Larus michahellis)

Asymmetry between males and females in the energy they invest initially in reproduction has resulted in the evolution of differing reproductive strategies (caring females vs. competitive males). However, parental care in many birds is shared by both sexes suggesting that male energy expenditure in agonistic behaviors and courtship feeding might compensate female costs of clutch production. Here, we tested the hypothesis that initial investment in reproduction by both sexes in the Yellow-legged Gull (Larus michahellis), a species with biparental care, is similar from a physiological perspective. In this income breeder, female and male reproductive investment during early breeding can be ultimately related to muscular activity (local foraging effort required for clutch production in females and courtship feeding and agonistic behaviors in the case of males). Thus, we evaluated sex-specific patterns of creatine kinase (CK, IU/L) levels in plasma, an indicator of physical effort associated with muscular activity dependent behaviors, through incubation as a reflection of the physiological response of both sexes to the reproductive investment they made up to clutch completion. Raw levels of CK were related to plasma levels of total proteins (TP, g/dL) to account for the differential physiological state of individuals when sampled (i.e. differential dehydratation). Female costs of clutch production were associated with post-laying levels of CK/TP. We grouped females according to their relative investment in clutch production: < 15.8%, 15.8 to 17.3% and > 17.3% of field metabolic rate; which showed increasing values of CK/TP (24.6, 53.1 and 66.0 IU/g, respectively). Moreover, we found similar CK/TP trends throughout incubation for both sexes (CK/TP = 50.2− [3.3 × days from laying]) suggesting similar physiological responses to reproductive effort and, therefore, analogous sex-specific initial investment. Thus, male investment in agonistic behaviors and courtship feeding apparently equaled female investment in clutch production. The use of CK measurements is revealed as a useful approach to investigating overall investment in reproduction for both sexes, providing further insights into our comprehension of reproductive strategies in seabirds.     

Sex-specific associations between reproductive output and hematozoan parasites of American kestrels

Parasites have the potential to decrease reproductive output of hosts by competing for nutrients or forcing hosts to invest in immune function. Conversely, reproductive output may affect parasite loads if hosts allocate resources to reproduction such that allocation to immune function is compromised. Both hypotheses implicitly have a temporal component, so we sampled parasites both before and after egg laying to examine the relationship between reproductive output (indexed using a combined measure of clutch size, egg volume, and initiation date) and blood parasite loads of American kestrels (Falco sparverius). Parasite loads measured prior to egg laying had no adverse effects on subsequent reproductive output. Females that previously had large reproductive outputs subsequently had lower parasite intensities than those whose outputs were smaller, suggesting that females were capable of allocating energy to both forming clutches and reducing parasite loads. Because male kestrels provide most of their mate's energetic needs before, during, and after egg laying, mate choice by females may have consequences for their parasite loads. Females choosing high-quality mates may not only have increased reproductive output, but may also obtain sufficient resources from their mates to enable them to reduce their parasite burdens. Males whose mates had large reproductive outputs were more likely to subsequently be parasitized and have more intense infections. For individual males sampled both before and after egg laying, those whose mates had larger reproductive outputs were also more likely to become parasitized, or remain parasitized, between sampling periods. Increased parasite loads of males may be one mechanism by which the costs of reproduction are paid.     

Life history of breeding partners alters age‐related changes of reproductive traits in a natural population of blue tits

Reproductive senescence, an intra‐individual decline in reproductive function with age, is widespread, but proximate factors determining its rate remain largely unknown. Most studies of reproductive senescence focus on females, leaving senescence in male function and its implications for female function largely understudied. We constructed linear mixed models to explore the interactive effects of paternal and maternal age and a life‐history trait (i.e. age at first reproduction) on four fitness components (i.e. laying date, clutch size, number of fledglings and number of recruits) measured in a wild, breeding population of blue tits Cyanistes caeruleus ogliastrae where individual breeding success has been followed for over 30 years (our dataset spanned 29 years). Previous studies have shown that, across female lifespan, laying date decreases and subsequently increases; earlier laying dates result in higher fitness because hatchlings have greater access to a seasonal food source. Our analyses reveal that females that initiate reproduction early in life show a greater delay in laying date with old age. In addition to delayed laying dates, older females lay smaller clutches. However, the magnitude of female age effects was influenced by the age at first reproduction of their breeding partners. Senescence of laying date and clutch size was reduced when females mated with males that reproduced early in life compared to males that delayed reproduction. We confirmed that both laying date and clutch size were significantly correlated with reproductive fitness suggesting that these dynamics early in the breeding cycle can have long‐term consequences. These complex phenotypic interactions shed light on the proximate mechanisms underlying reproductive senescence in nature and highlight the potential importance of cross‐sex age by life‐history interactions.     

Reproductive effort decreases antibody responsiveness

The prevalence and intensity of parasitic infection often increases in animals when they are reproducing. This may be a consequence of increased rates of parasite transmission due to reproductive effort. Alternatively, endocrine changes associated with reproduction can lead to immunosuppression. Here we provide support for a third potential mechanism: reduced immunocompetence as a consequence of adaptive reallocation of resources in times of increased energetic demand. In captive zebra finches Taeniopygia guttata, reproductive effort was manipulated through brood size. Enhanced effort was found to affect the production of antibodies towards sheep red blood cells. In addition, activity of zebra finches was manipulated independently of parental care. Experimentally increased daily workloads in activity reward schedules also suppressed antibody production. Thus, we show that not just the reproductive state, but the increased activity that accompanies reproduction is associated with immunocompetence. This mechanism may be sufficient to explain the increased parasitism observed in reproducing animals. We suggest that reduced immunocompetence as a consequence of increased reproductive effort may be an important pathway for the life history cost of reproduction.     

Reproductive effort reduces specific immune response and parasite resistance

If a trade-off exists between reproductive effort and immune function, life-history decisions may have important implications for parasite resistance. Here, we report effects of experimental manipulation of reproductive effort on subsequent specific immune function and parasite resistance in the collared flycatcher, Ficedula albicollis. Our results show that increased reproductive effort of females immunized with Newcastle disease virus (NDV) vaccine negatively affected the ability to respond with NDV-specific antibodies. We further show that increased reproductive effort increased the intensity of Haemoproteus infections and that such infections are associated with higher mortality. Our results thus provide support for the hypothesis that immune suppression caused by reproductive effort may be an important mechanism mediating the life-history cost of reproduction.     

Gender-specific costs of reproduction on vegetative growth and physiological performance in the dioecious shrub Corema album

Background and Aims

Reproductive costs imply trade-offs in resource distribution at the physiological level, expressed as changes in future growth and/or reproduction. In dioecious species, females generally endure higher reproductive effort, although this is not necessarily expressed through higher somatic costs, as compensatory mechanisms may foster resource uptake during reproduction.

Methods

To assess effects of reproductive allocation on vegetative growth and physiological response in terms of costs and compensation mechanisms, a manipulative experiment of inflorescence bud removal was carried out in the sexually dimorphic species Corema album. Over two consecutive growing seasons, vegetative growth patterns, water status and photochemical efficiency were measured to evaluate gender-related differences.

Key Results

Suppression of reproductive allocation resulted in a direct reduction in somatic costs of reproduction, expressed through changes in growth variables and plant physiological status. Inflorescence bud removal was related to an increase in shoot elongation and water potential in male and female plants. The response to inflorescence bud removal showed gender-related differences that were related to the moment of maximum reproductive effort in each sexual form: flowering in males and fruiting in females. Delayed costs of reproduction were found in both water status and growth variables, showing gender-related differences in resource storage and use.

Conclusions

Results are consistent with the existence of a trade-off between reproductive and vegetative biomass, indicating that reproduction and growth depend on the same resource pool. Gender-related morphological and physiological differences arise as a response to different reproductive resource requirements. Delayed somatic costs provide evidence of gender-related differences in resource allocation and storage. Adaptive differences between genders in C. album may arise through the development of mechanisms which compensate for the cost of reproduction.     

Allocation from capital and income sources to reproduction shift from first to second clutch in the flesh fly, Sarcophaga crassipalpis

Understanding how resources are allocated between survival and reproduction is fundamental to the study of the evolution of life histories. Reproductive resources can come from two intrinsic resource pools, stored reserves (capital) acquired before reproduction or income acquired during reproduction. The variety of reproductive strategies in insects is remarkable and reproductive allocation encompasses the complete range of allocation strategies from pure capital breeders to pure income breeders. However, most organisms probably use a blend of capital and income and this blend is likely dynamic, changing between reproductive bouts in response to internal and external conditions. We used stable isotopes to quantify the allocation of capital and income resources to reproduction in the flesh fly, Sarcopha crassipalpis and assessed how allocation patterns change over multiple bouts of reproduction. Sarcophaga crassipalpis shifts from a slight investment of capital in the first clutch to an almost pure income breeder in the second clutch. We discuss the relationship between activity and allocation, and the potential for this system to understand how allocation patterns change in response to environmental stress.     

Sex differences in the effects of juvenile and adult diet on age‐dependent reproductive effort

Sexual selection should cause sex differences in patterns of resource allocation. When current and future reproductive effort trade off, variation in resource acquisition might further cause sex differences in age‐dependent investment, or in sensitivity to changes in resource availability over time. However, the nature and prevalence of sex differences in age‐dependent investment remain unclear. We manipulated resource acquisition at juvenile and adult stages in decorated crickets, Gryllodes sigillatus, and assessed effects on sex‐specific allocation to age‐dependent reproductive effort (calling in males, fecundity in females) and longevity. We predicted that the resource and time demands of egg production would result in relatively consistent female strategies across treatments, whereas male investment should depend sharply on diet. Contrary to expectations, female age‐dependent reproductive effort diverged substantially across treatments, with resource‐limited females showing much lower and later investment in reproduction; the highest fecundity was associated with intermediate lifespans. In contrast, long‐lived males always signalled more than short‐lived males, and male age‐dependent reproductive effort did not depend on diet. We found consistently positive covariance between male reproductive effort and lifespan, whereas diet altered this covariance in females, revealing sex differences in the benefits of allocation to longevity. Our results support sex‐specific selection on allocation patterns, but also suggest a simpler alternative: males may use social feedback to make allocation decisions and preferentially store resources as energetic reserves in its absence. Increased calling effort with age therefore could be caused by gradual resource accumulation, heightened mortality risk over time, and a lack of feedback from available mates.     

A pre‐breeding immune challenge delays reproduction in the female dark‐eyed junco Junco hyemalis

Precise timing of life‐history transitions in predictably changing environments is hypothesized to aid in individual survival and reproductive success, by appropriately matching an animal's physiology and behavior with prevailing environmental conditions. Therefore, it is imperative for individuals to time energetically costly life‐history stages (i.e. reproduction) so they overlap with seasonal peaks in food abundance and quality. Female lifetime reproductive fitness is affected by several factors that influence energy balance, including arrival date, timing of egg production, and energetic condition. Therefore, any extra energetic costs during reproduction may negatively affect timing of egg production, and ultimately a female's fitness. For example, mounting an immunological response elicits a high energetic cost, and this transfer of resources towards cell and immune system maintenance could have direct negative effects on reproductive timing. In order to determine whether an immune challenge delays onset of breeding (i.e. egg production), we administered either a humoral immune challenge (keyhole limpet hemocyanin (KLH)) (treatment) or physiological saline (control) to free‐living female dark‐eyed juncos Junco hyemalis in the period immediately prior to egg‐laying (～4 weeks). We found that KLH‐injected females artificially delayed clutch initiation when compared to control females. These data help to refine our understanding of how free‐living birds allocate resources between reproduction and self‐maintenance processes during the critical pre‐laying period of the annual cycle.     

Resource use for reproduction depends on spring arrival time and wintering area in an arctic breeding shorebird

Resources for egg production may come from body reserves stored before breeding (“capital breeders”) or from food acquired at the breeding site (“income breeders”). Arctic migrants were long thought to be capital breeders, because they often arrive at a time when local food availability is still limited. However, later evidence suggested that arctic breeding shorebirds are primarily income breeders, or that they use a mixed strategy depending on laying date. We explored the relationship between laying date and resource use for reproduction in the pectoral sandpiper Calidris melanotos breeding in the Alaskan arctic by contrasting carbon isotope (δ¹³C) values of the local diet and of maternal plasma, cellular blood, feather and claw with those of the eggs produced. Our results revealed that early breeding females utilize resources for egg production that were acquired recently at staging areas, whereas later breeding females mostly relied on nutrients derived from local food sources. These findings suggest that the resource allocation strategy used for reproduction differs among females, and varies depending on the timing of arrival and the start of reproduction. The arrival date at the breeding ground and laying date may critically depend on non‐breeding season events such as winter habitat choice, staging areas or migration routes. By comparing maternal feather δ¹³C, claw δ¹³C and feather δD, we examined whether non‐breeding season events influenced the use of resources for egg production through variation in capture date or clutch initiation date. Female pectoral sandpipers originating from moulting areas characterized by higher (more positive) δD signatures were caught earlier and started laying earlier, and they used stored resources for reproduction. Using regional maps of δD values for precipitation in the wintering sites in South America, we compared the spatial variation in the observed feather δD signatures. This analysis indicated that female pectoral sandpipers with higher δD signatures, presumably coming from more north‐easterly wintering sites in southern America, started laying earlier and used mostly stored resources for egg production, compared to females that wintered (or at least moulted) further south. Our results thus show that winter moulting habitat is linked to breeding resource allocation strategy in this high‐arctic breeding shorebird.     

Costs of incubation and immunocompetence in the collared flycatcher

This paper investigates the costs of incubation in terms of reduced reproductive success and investigates whether incubation competes with immune function for resources. I performed a clutch size manipulation experiment in which two eggs were either removed from or added to the nests of collared flycatchers, Ficedula albicollis, for 1 week during incubation and subsequently returned to their original nests before hatching. To induce immune response, the females were challenged with sheep red blood cells. While the duration of incubation, hatching success and fledgling number did not differ between experimental groups, fledgling condition was significantly lower in broods that had been enlarged during incubation. Neither the females' condition nor their ability to respond to a novel antigen differed between treatments. The relationship between antibody production and female condition was significantly positive, but only among females incubating reduced clutches. I conclude that the costs of incubation in the collared flycatcher are not negligible and are manifested only at the chick-rearing phase.     

COSTS OF REPRODUCTION IN THE WILD: PATH ANALYSIS OF NATURAL SELECTION AND EXPERIMENTAL TESTS OF CAUSATION

During 1991 through 1994, natural selection on reproductive effort in side-blotched lizards was indexed by measuring total clutch mass produced on the first clutch of the reproductive season and assessing how such effort in current reproduction affects subsequent survival and clutch production. In addition, selection was also experimentally assessed in free-ranging female side-blotched lizards by (1) surgically decreasing total clutch mass (direct ovarian manipulation) and enhancing clutch mass using (2) exogenous gonadotropin, and (3) exogenous corticosterone. Surgical reduction of clutch mass uniformly enhanced survival. However, increasing clutch mass had more complex effects depending on year. Experimentally enhanced clutch mass enhanced survival in 1991, had no effect on survival in 1992, and decreased survival in 1993. Despite the complexity of these experimental results, they are corroborated by our comparative data. It is important to note that local environmental effects can obscure detection of costs arising from natural variation in reproductive effort, and we removed such effects using path analysis. The striking shift in natural selection favoring females laying a large clutch mass (1991) to selection against females laying a large clutch mass (1993) is associated with an end of a severe multiyear drought. Our natural-history observations suggest that the correlated increase in predatory snake activity on our study site, coincident with the end of the drought, is the agent of natural selection. Although the actual agents of selection (e.g., snake predation versus drought-related effects) are not resolved, the patterns of natural selection measured in our comparative and experimental data are also consistent with year-to-year changes in clutch mass and egg size that would be indicative of rapid short-term evolution in these traits.     

Reproductive investment is connected to innate immunity in a long-lived animal

Life-history theory predicts that organisms optimize their resource allocation strategy to maximize lifetime reproductive success. Individuals can flexibly reallocate resources depending on their life-history stage, and environmental and physiological factors, which lead to variable life-history strategies even within species. Physiological trade-offs between immunity and reproduction are particularly relevant for long-lived species that need to balance current reproduction against future survival and reproduction, but their underlying mechanisms are poorly understood. A major unresolved issue is whether the first-line innate immune function is suppressed by reproductive investment. In this paper, we tested if reproductive investment is associated with the suppression of innate immunity, and how this potential trade-off is resolved depending on physiological state and residual reproductive value. We used long-lived capital-breeding female eiders (Somateria mollissima) as a model. We showed that the innate immune response, measured by plasma bacteria-killing capacity (BKC), was negatively associated with increasing reproductive investment, i.e., with increasing clutch size and advancing incubation stage. Females in a better physiological state, as indexed by low heterophil-to-lymphocyte (H/L) ratios, showed higher BKC during early incubation, but this capacity decreased as incubation progressed, whereas females in poorer state showed low BKC capacity throughout incubation. Although plasma BKC generally declined with increasing H/L ratios, this decrease was most pronounced in young females. Our results demonstrate that reproductive investment can suppress constitutive first-line immune defence in a long-lived bird, but the degree of immunosuppression depends on physiological state and age.     

Nest-building activity and laying date influence female reproductive investment in magpies: an experimental study

Nest size or nest-building activity has recently been hypothesized to be a postmating sexually selected signal in monogamous birds: females may assess a male's parental quality and willingness to invest in reproduction by his participation in nest building. Females may thus adjust their reproductive effort (i.e. clutch size) not only to their own abilities but also to those of their mates. We investigated whether female magpies, Pica pica, use nest-building activity rather than nest size to adjust their reproductive effort during replacement breeding attempts. After we removed their first clutch, high-quality pairs that built a large nest for the first clutch were more capable of building a replacement nest and females adjusted their clutch size in relation to the time it took to build the nest rather than nest size. We also found support for the hypothesized trade-off between clutch size and egg size in magpies. In replacement clutches females decreased clutch size and increased egg volume, thereby probably improving the survival probability of their offspring in less favourable conditions.Copyright 2002 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour     

Variation in reproductive output of marine turtles

Most marine turtle species are non-annual breeders and show variation in both the number of eggs laid per clutch and the number of clutches laid in a season. Large levels of inter-annual variation in the number of nesting females have been well documented in green turtle nesting populations and may be linked to environmental conditions. Other species of marine turtle exhibit less variation in nesting numbers. This inter-specific difference is thought to be linked to trophic status. To examine whether individual reproductive output is more variable in the herbivorous green turtle (Chelonia mydas Linneaeus 1758) than the carnivorous loggerhead (Caretta caretta Linneaeus 1758), we examined the nesting of both species in Cyprus over nine seasons. Green turtles showed slower annual growth rates (0.11 cm year⁻¹ curved carapace length (CCL) and 0.27 cm year⁻¹ curved carapace width (CCW)) than loggerhead turtles (0.36 cm year⁻¹ CCL, 0.51 cm year⁻¹ CCW). CCL was highly correlated to mean clutch size in both green (R²=0.51) and loggerhead turtles (R²=0.61) and maximal clutch size of green turtles (R²=0.58). Larger females did not lay a greater number of clutches or have a shorter remigration interval than smaller females of either species. On average, the size of green turtle clutches increased and that of loggerhead turtles decreased as the season progressed. Individual green turtles, however, produced more eggs per clutch through the season to a maximum in the third or fourth clutch. In loggerhead turtles, clutches 1-4 were very similar in size but the fifth clutch was 38% smaller than the first. No individuals of either species were recorded laying more than five clutches. Green turtles may not be able to achieve their maximum reproductive output with respect to clutch size throughout the season, whereas only loggerhead turtles laying five clutches (n=5) appear to become resource depleted. Green turtles nesting in years when large numbers of nests were recorded laid a greater number of clutches than females nesting in years with lower levels of nesting.     

Sex-specific response to nutrient limitation and its effects on female mating success in a gift-giving butterfly

Animals with complex life cycles respond to early food limitation by altering the way resources are allocated in the adult stage. Response to food limitation should differ between males and females, especially in organisms whose mating systems include nutritional nuptial gifts. In these organisms, males are predicted to keep their allocation to reproduction (sperm and nuptial gift production) constant, while females are predicted to sacrifice allocation to reproduction (egg production) since they can compensate by acquiring nuptial gifts when mating. In this study, we investigated how dietary nitrogen limitation during the larval stage affects sex-specific resource allocation in Pieris rapae butterflies. Also, we tested whether nutrient-limited females increased nuptial gift acquisition as a way to compensate for low allocation to reproduction. We found that as predicted females, but not males, sacrifice allocation to reproduction when larval dietary nitrogen is limited. However, females were unable to compensate for this low reproductive allocation by increasing their mating rate to acquire additional gifts. Females reared on low nitrogen diets also reduced wing coloration, a potential signal of female fecundity status. We suggest that female mating frequency is constrained by male mate choice based on females’ wing coloration. This study provides new insights into how larval dietary nitrogen, a key nutritional resource for all herbivores, alters male and female allocation to reproduction as well as to ornamentation.     

Life-history evolution in Anodonta piscinalis (Mollusca,Pelecypoda)

Summary The correlations between certain life-history parameters (reproductive effort, reproductive life-span, age of first reproduction, general growth index, variation in juvenile survival, availability of resources) were studied in 13 populations of the mussel Anodonta piscinalis in south-western Finland in 1975 and 1976Reproductive life-span correlated positively (r _s=0,823, P<0.001) with variation in juvenile survival. The average availability of resources correlated negatively both with the reproductive life-span (r _s=-0.841, P<0.001 after the variation by juvenile survival had been deleted) and variation in juvenile survival (r _s=-0.676, P<0.05).The reproductive effort for female mussels at each site was computed by comparing body weight of reproductive females with body weights of non-reproductive individuals. Availability of resources was much higher in 1976 than in 1975. Consequently, the reproductive effort, an index of the strain of reproduction, was higher in 1975 than in 1976. In 1975 there was a significant correlation between reproductive effort and the length of the reproductive life-span (-0.727, -0.806) and also with the reproductive effort and the variation in juvenile survival (-0.718, -0.758) in females of the length of 60 mm and 70 mm respectively. In 1976, when availability of resources was better, such correlations were not found.Spatial and temporal change in the intake of resources complicates applicability of the principle of resource allocation in the theory of life-history evolution. Studying the mere allocation is not enough if the intake of resources varies in the groups studied. The ratio ovary weight/body weight is a dubious measure of reproductive effort in comparative studies when the input of resources can vary, and this possibility can be ruled out only exceptionally.Correlation between growth and reproductive effort was positive, obviously because both are important components in creating high reproductive capacity. In 1975 reproductive effort increased with size (age). Change in reproductive effort correlated with reproductive life-span (r _s=-0.633, P<0.05).The following parameters occurred together: short reproductive life-span, low age of first reproduction, high reproductive effort, rapid growth and high clutch size. They were realized at sites where the availability of resources was good and variation in juvenile survival was low-i.e. the environment was stable. The results conflict with the prediction of the theory of r and K-selection.