Offspring growth, survival and reproductive success in the bank vole: a litter size manipulation experiment

To estimate the optimality of brood size, it is essential to study the effects of brood size manipulation on offspring survival and reproductive success. Moreover, testing the generality of the hypothesis of reproductive costs requires experimental data from a diversity of organisms. Here I present data on the growth, survival and reproductive success of bank vole Clethrionomys glareolus individuals from manipulated litters. Furthermore, the survival of mothers whose litter size was manipulated was studied. At weaning, the mean weight of pups from enlarged litters was lower and from reduced litters higher compared to control litters. After winter, at the start of the breeding season, individuals from enlarged litters, especially males, were still lighter than individuals from the other two treatments. Litter enlargements did not increase the number of reproducing female offspring per mother, nor did the litter sizes of female offspring differ between treatments. There were no differences between treatments in winter survival of offspring after weaning, but among female offspring, weaning weight explained the survival probabilities over winter. A higher weight of females at winter determined the probability of starting to reproduce in spring. The survival of mothers did not seem to be influenced by litter manipulation performed the previous year. According to the results, mothers nursing enlarged or reduced litters do not gain any fitness benefits in terms of number of offspring surviving to breeding. The results are consistent with the majority of experiments conducted in birds, which have found costs of enlarged brood appearing as offspring trade-offs rather than parent trade-offs. Received: 14 December 1997 / Accepted: 1 March 1998     

Limitation of reproductive success by food availability and litter size in the bank vole,Clethrionomys glareolus

Food limitation has been suggested as one of the most important factors affecting life history evolution in terrestrial vertebrates. However, this inference is based mainly on evidence from birds, and reproductive trade-offs may differ among groups with different forms of parental care. To study whether the costs of enlarged litters (decreased mass of offspring) would appear when food is not limiting, we performed outdoor enclosure experiments in which we manipulated simultaneously the litter size (control versus litter plus two pups) and food availability (control versus food-supplemented) of female bank voles, Clethrionomys glareolus. The weaning success of females increased significantly in response to supplementary food. When females were provided with extra food, no differences were observed in body masses of weanlings of control and enlarged litters. Further, food-supplemented females grew to larger sizes during nursing than unsupplemented females. Our experiment suggests that energetic requirements during nursing constrain the number of offspring that can be raised successfully in a particular breeding attempt. The results also indicate that unlimited food resources may increase future reproductive potential of females because they can use more energy for somatic growth.     

Infanticide in the evolution of reproductive synchrony: effects on reproductive success

Synchronous breeding in animals and plants has stimulated both a theoretical and empirical examination of the possible benefits of active synchronization. The selective pressures of predation and infanticide are the strongest candidates proposed to explain the evolution of reproductive synchrony. Alternatively, breeding asynchronously with conspecifics may ensure a greater availability of resources per breeder. However, the possible fitness benefits resulting from active asynchronization have not yet received attention in evolutionary ecology. Here we present a hypothesis, based on a graphical model, illustrating the costs and benefits of the two modes of reproduction. We tested the hypothesis empirically using a 2 x 2 full factorial study design, where reproductive synchrony and infanticide tactics were manipulated in enclosed populations of the bank vole. The results reveal a relationship between infanticide tactics and breeding synchrony as illustrated by our hypothesis. In general, female reproductive success (number and size of offspring surviving to weaning) was significantly lower in infanticidal populations. Moreover, an asynchronous breeding pattern proved to be advantageous in the noninfanticidal population but this advantage of asynchrony was lost as infanticide became common in the population. Our findings support the idea that synchronous reproduction could have evolved as a counterstrategy against infanticide.     

Determinants of reproductive success in voles: space use in relation to food and litter size manipulation

Spacing behaviour of female mammals is suggested to depend on the distribution and abundance of food. In addition, food limitation has been found to constrain the reproductive success of females. However, whether females maximize their reproductive success by adjusting space use in relation to current food availability and reproductive effort (e.g. litter size) has not been experimentally studied. We examined these questions by manipulating simultaneously food resources (control vs. food supplementation) and litter sizes (control vs. plus two pups) of territorial female bank voles (Clethrionomys glareolus) in large outdoor enclosures. Females with supplementary food had smaller home ranges (foraging area) and home range overlaps than control females, whereas litter size manipulation had no effect on space use. In contrast, the size of territory (exclusive area) was not affected by food supplementation or litter size manipulation. As we have previously shown elsewhere, extra food increases the reproductive success of bank vole females in terms of size and proportion of weaned offspring. According to the present data, greater overlap of female home ranges had a negative effect on reproductive success of females, particularly on survival of offspring. We conclude that higher food availability increases the reproductive success of bank vole females, and this effect may be mediated through lower vulnerability of offspring to direct killing and/or detrimental effects from other females in the population. Moreover, it seems that when density of conspecifics is controlled for, home range sizes of females, but not territoriality, is related to food resources in Clethrionomys voles.     

Optimal allocation of reproductive effort: manipulation of offspring number and size in the bank vole

The number of offspring attaining reproductive age is an important measure of an individual's fitness. However, reproductive success is generally constrained by a trade-off between offspring number and quality. We conducted a factorial experiment in order to study the effects of an artificial enlargement of offspring number and size on the reproductive success of female bank voles (Clethrionomys glareolus). We also studied the effects of the manipulations on growth, survival and reproductive success of the offspring. Potentially confounding effects of varying maternal quality were avoided by cross-fostering. Our results showed that the number of offspring alive in the next breeding season was higher in offspring number manipulation groups, despite their smaller body size at weaning. Offspring size manipulation had no effect on offspring growth or survival. Further, the first litter size of female offspring did not differ between treatments. In conclusion, females may be able to increase the number of offspring reaching reproductive age by producing larger litters, whereas increasing offspring size benefits neither the mother nor the offspring.     

Reproductive effort of female bank voles in a risky environment

In cyclically fluctuating vole populations, strong intraspecific competition and intense predation simultaneously or separately increase the costs of reproduction and so may set the framework for the optimal breeding tactic of voles. In a factorial experiment, we manipulated two factors in the breeding environment of bank vole (Clethrionomys glareolus) pairs, each with two treatment levels: no predation risk or high risk of specialist predators and low or high density of voles. In the manipulation, we used odours of conspecific voles and/or mustelid predators. Both over-wintered and young, summer-born, wild-caught bank vole females and males in breeding condition were used. Each female—male pair of voles was placed in a cage and the cages were distributed in large outdoor enclosures. All animals were fed ad libitum. Under predation risk, both old and young females suppressed breeding significantly. The density of conspecific voles did not affect overall breeding. However, there was some evidence that population density stimulated breeding of old females but suppressed breeding of young ones. Both risk factors appeared to increase litter sizes of those individuals who decided to breed. Our results indicate that the risk of predation may be an important factor determining reproductive tactics of bank vole females. In risky environments, females seemed to choose between two totally opposite tactics: they suppressed breeding, which may increase their own survival to the next breeding event, or they continued to breed in spite of expected high survival costs. Females seemed to compensate the latter costs with a higher effort to the current and probably the last reproduction.     

Contrasting effects of large density changes on relative testes size in fluctuating populations of sympatric vole species

Across species, there is usually a positive relationship between sperm competition level and male reproductive effort on ejaculates, typically measured using relative testes size (RTS). Within populations, demographic and ecological processes may drastically alter the level of sperm competition and thus, potentially affect the evolution of testes size. Here, we use longitudinal records (across 38 years) from wild sympatric Fennoscandian populations of five species of voles to investigate whether RTS responds to natural fluctuations in population density, i.e. variation in sperm competition risk. We show that for some species RTS increases with density. However, our results also show that this relationship can be reversed in populations with large-scale between-year differences in density. Multiple mechanisms are suggested to explain the negative RTS–density relationship, including testes size response to density-dependent species interactions, an evolutionary response to sperm competition levels that is lagged when density fluctuations are over a certain threshold, or differing investment in pre- and post-copulatory competition at different densities. The results emphasize that our understanding of sperm competition in fluctuating environments is still very limited.     

Genetic estimates of annual reproductive success in male brown bears: the effects of body size, age, internal relatedness and population density

1. We studied male yearly reproductive success (YRS) and its determinants (phenotypic characteristics, age, population density) in two Scandinavian brown bear populations, using molecular techniques to determine paternity. 2. We found a significant difference in male YRS between the study areas, with lower YRS in the south than in the north. 3. In general, older and larger males had higher YRS. Older males may be more experienced in competition for reproduction (male dominance). Large body size is of direct benefit in male-male competition and of advantage in endurance competition for the access to females. 4. Age was relatively more important for YRS in the north and body size was more important in the south, due perhaps to differences in male age structure due to illegal killing. A single old male dominated the reproduction in the north during the study, which resulted most probably in the relatively higher importance of age in the north. In the south, with a more even male age structure, no single male was able to dominate, probably resulting in a more intense competition among males, with body size as the deciding factor. 5. Male YRS was correlated positively with population density. This may be related to the structure of the expanding bear population, with female densities declining towards the population edge. 6. Internal relatedness, a measure of genetic heterozygosity, was correlated negatively with YRS, suggesting that outbred individuals have a higher YRS. Individual heterozygosity at key or many loci may reflect male physical qualities and condition-sensitive traits, which may benefit males directly in contest or in sperm competition.     

Allometry of territory size and metabolic rate as predictors of self-thinning in young-of-the-year Atlantic salmon

1. Self-thinning is a progressive decline in population density caused by competitively induced losses in a cohort of growing individuals and can be depicted as: log₁₀ (density) = c − β log₁₀ (body mass).
2. In mobile animals, two mechanisms for self-thinning have been proposed: (i) the space hypothesis predicts that maximum population density for a given body size is the inverse of territory size, and hence, the self-thinning slope is the negative of the slope of the allometric territory-size relationship; (ii) the energetic equivalence hypothesis predicts that the self-thinning slope is the negative of the slope of the allometric metabolic rate relationship, assuming a constant supply of energy for the cohort.
3. Both hypotheses were tested by monitoring body size, population density, food availability and habitat for young-of-the-year Atlantic salmon ( Salmo salar ) in Catamaran Brook, New Brunswick. The results were consistent with the predictions of the space hypothesis. Observed densities did not exceed the maximum densities predicted and the observed self-thinning slope of −1·16 was not significantly different from the slope of −1·12, predicted by the allometry of territory size for the population under study.
4. The observed self-thinning slope was significantly steeper than −0·87, predicted by the allometry of metabolic rate, perhaps because of a gradual decline in food abundance over the study period. The decline in density was more rapid in very shallow sites and may have been partly caused by a seasonal change in water depth and an ontogenetic habitat shift rather than solely by competition for food or space.
5. The allometry of territory size may be a useful predictor of self-thinning in populations of mobile animals competing for food and space.     

Assessing the impact of density dependence in field populations of Aedes aegypti

Although many laboratory studies of intra-specific competition have been conducted with Ae. aegypti, there have been few studies in natural environments and none that examined density dependence in natural containers at normal field densities. Additionally, current mathematical models that predict Ae. aegypti population dynamics lack empirically-based functions for density-dependence. We performed field experiments in Tapachula, Mexico, where dengue is a significant public health concern. Twenty-one containers with natural food and water that already contained larvae were collected from local houses. Each container was divided in half and the naturally occurring larvae were apportioned in a manner that resulted in one side of the container (high density) having four times the density of the second side (low density). Larvae were counted and pupae were removed daily. Once adults emerged, wing span was measured to estimate body size. Density had a significant impact on larval survival, adult body size, and the time taken to transition from 4(th) instar to pupation. Increased density decreased larval survival by 20% and decreased wing length by an average of 0.19 mm. These results provide a starting point for a better understanding of density dependence in field populations of Ae. aegypti.     

Size variation and reproductive success in the mosquito Aedes cantans

Abstract. A link between density-dependent larval competition and adult size of the mosquito Aedes cantans was demonstrated in northern England. Ponds containing high larval densities produced smaller larvae which, in turn, resulted in smaller adults at emergence.
In both 1989 and 1990, parous mosquitoes caught at human bait were larger than nulliparous ones, suggesting that larger mosquitoes are more successful at host location and egg-laying and also that they may be longer lived.
Larger mosquitoes produced larger egg clutches than smaller females: however, there was no difference in the size of eggs laid by large and small females.     

杭州石荠苎(Mosla hangchowensis)种群密度制约实验的统计分析

本文研究了我国特有、分布区极狭窄的一年生草本植物——杭州石荠苎(Mosla hangchowensis)种群的密度制约规律。结果表明：在生长季内,种群的死亡率与密度密切相关。种群的最适密度为200～1000株/m²左右。不同密度种群的平均株高、开花数等性状随时间的动态关系均符合“logistic”模型。高密度种群中60%左右的个体能完成生活史;低密度种群中80%以上的个体能完成生活史。种群密度较高制约杭州石荠苎的植株形态和繁殖投资。     

An Experimental study of reduced parental effort and future reproductive success in the puffin, Fratercula arctica

1. The puffin, a long-lived seabird, was studied on the Isle of May, East Scotland between 1990 and 1992. During two of these years, parental effort was experimentally decreased by supplementary feeding of young. This aimed to identify inter-year reproductive costs, and show whether they took the form of reduced adult survival, reduced fledging success and/or a reduction in the 'quality' of offspring in the following year.
2. The feeding treatment significantly reduced the daily number of feeds delivered by experimental parents by 67% in 1990 and 87% in 1991.
3. The proportions of experimental and control parents returning to the colony in the year following manipulation did not differ significantly, although in 1991, 2·5 times as many controls (young unfed) as experimental birds (young fed) failed to return.
4. The fledging success of experimental pairs in the year following manipulation (68%) was significantly higher than that of controls (24%).
5. Experimental pairs raised young with significantly higher body condition (Residual Peak Mass) than that of controls in the year following manipulation (1992).
6. Experimental parents did not differ from controls in their body condition (Lipid Reserve Mass) or rate of reserve depletion, either in the year of manipulation or in the following breeding season; hence there was no evidence for a role of the measured component of body condition in the cost mechanism.
7. The study demonstrated inter-year reproductive costs for puffins and supported the hypothesis that long-lived species reduce the 'quality' of their offspring or abandon a breeding attempt rather than compromise their survival and future opportunities to reproduce.     

Climatic effects on the breeding phenology and reproductive success of an arctic-nesting goose species

Climate warming is pronounced in the Arctic and migratory birds are expected to be among the most affected species. We examined the effects of local and regional climatic variations on the breeding phenology and reproductive success of greater snow geese ( Chen caerulescens atlantica ), a migratory species nesting in the Canadian Arctic. We used a long-term dataset based on the monitoring of 5447 nests and the measurements of 19 234 goslings over 16 years (1989–2004) on Bylot Island. About 50% of variation in the reproductive phenology of individuals was explained by spring climatic factors. High mean temperatures and, to a lesser extent, low snow cover in spring were associated with an increase in nest density and early egg-laying and hatching dates. High temperature in spring and high early summer rainfall were positively related to nesting success. These effects may result from a reduction in egg predation rate when the density of nesting geese is high and when increased water availability allows females to stay close to their nest during incubation recesses. Summer brood loss and production of young at the end of the summer increased when values of the summer Arctic Oscillation (AO) index were either very positive (low temperatures) or very negative (high temperatures), indicating that these components of the breeding success were most influenced by the regional summer climate. Gosling mass and size near fledging were reduced in years with high spring temperatures. This effect is likely due to a reduced availability of high quality food in years with early spring, either due to food depletion resulting from high brood density or a mismatch between hatching date of goslings and the timing of the peak of plant quality. Our analysis suggests that climate warming should advance the reproductive phenology of geese, but that high spring temperatures and extreme values of the summer AO index may decrease their reproductive success up to fledging.     

Costs of mating competition limit male lifetime breeding success in polygynous mammals

Although differences in breeding lifespan are an important source of variation in male fitness, the factors affecting the breeding tenure of males have seldom been explored. Here, we use cross-species comparisons to investigate the correlates of breeding lifespan in male mammals. Our results show that male breeding lifespan depends on the extent of polygyny, which reflects the relative intensity of competition for access to females. Males have relatively short breeding tenure in species where individuals have the potential to monopolize mating with multiple females, and longer ones where individuals defend one female at a time. Male breeding tenure is also shorter in species in which females breed frequently than in those where females breed less frequently, suggesting that the costs of guarding females may contribute to limiting tenure length. As a consequence of these relationships, estimates of skew in male breeding success within seasons overestimate skew calculated across the lifetime and, in several polygynous species, variance in lifetime breeding success is not substantially higher in males than in females.     

Matrilineal kin clusters and their effect on reproductive success in the field vole Microtus agrestis

Kin dusters of philopatric females are thought to form the basicsocial organization of breeding females in various vole species.The spatial association between young breeding females and theirmothers was studied in five 2500–m² enclosures in naturalhabitats during mid- and late breeding season in summer 1992.The mothers (founder females) were introduced into the enclosuresin die beginning of the study in June, whereas the young breedingfemales were born within the enclosures. Matrilineal kinshipwas determined by using radionudides. In July-August, 58% ofyoung breeding females were philopatric, but 82% of the mother-daughterassociations broke up until August-September. Of the young femalesthat started to breed in August-September, only 12% were philopatricThus the kin clusters found were short Hved and few in numberduring the late breeding season. The reduction in the incidenceof the kin clusters coincided with an increase of populationdensity. The mean reproductive success of females breeding inkin clusters was better than that of those breeding separately.In addition, among the nonphilopatric females there was a positive,significant correlation between reproductive success and thedistance to the nearest unrelated founder female. Such correlationwas not found among the philopatric females.