Determinants and consequences of age of primiparity in bighorn ewes

Because variation in age of first reproduction can have major effects on individual fitness and population dynamics, it is important to understand what maintains that variability. Although early primiparity is assumed to be costly, it is sometimes associated with high lifetime reproductive success. We used a long‐term study on bighorn sheep Ovis canadensis to determine what variables affect age at first reproduction, investigate the impact of primiparity on body resources and quantify the reproductive performance of primiparous ewes. We then examined the consequences of delayed primiparity on adult body mass, longevity and lifetime reproductive success. Environmental conditions during early development, body mass as a yearling, genotype and maternal effects affected age of primiparity. Primiparous ewes lost more mass in winter and gained less mass in summer than multiparous ewes. Small yearling ewes that postponed reproduction attained similar adult mass than heavy yearling ewes who reproduced at a younger age. Early primiparity did not reduce longevity and was positively associated with lifetime reproductive success. Starting to reproduce as soon as possible appears to maximize fitness of females. When early life conditions are unfavorable, however, delayed primiparity allows greater body growth and likely maximizes survival. The combination of a conservative reproductive strategy and maternal effects on age of primiparity may partly delay population recovery following density‐dependent declines.     

Mass- and density-dependent reproductive success and reproductive costs in a capital breeder

For capital breeders, mass may affect reproductive potential. Reproductive expenditure may reduce future reproductive potential, particularly when resources are scarce. To test the hypothesis that reproductive success and the costs of reproduction vary according to mass and population density, we analyzed 25 yr of data on bighorn ewes (Ovis canadensis). The number of adult females was first limited by yearly removals, then allowed to triple. We found no survival costs of reproduction for ewes aged 4-7 yr. For ewes aged 8-14 yr, survival was density dependent for barren ewes but not for ewes that weaned lambs. Failure to lamb was rare and negatively correlated with fertility the following year. At low population density, lactation had a negative effect on mass gain but had a limited reproductive cost. At high density, heavy ewes had higher reproductive success than light ewes, and the reproductive cost and somatic costs of reproduction increased. The cost of reproduction was greater for light than for heavy ewes. Survival of weaned lambs to 1 yr was affected by population density but not by maternal mass or previous reproductive success. In large mammals, manipulations of reproductive effort are problematic, but long-term monitoring of individual mass and reproductive success under varying conditions of resource availability can provide insights into the evolution of life histories.     

Compensatory Growth Limits Opportunities for Artificial Selection in Alpine Chamois

ABSTRACT In ungulates, big males with large weapons typically outcompete other males over access to estrous females. In many species, rapid early growth leads to large adult mass and weapon size. We compared males in one hunted and one protected population of Alpine chamois (Rupicapra rupicapra) to examine the relationship between horn length and body mass. We assessed whether early development and hunter selectivity affected age-specific patterns of body and horn size and whether sport hunting could be an artificial selection pressure favoring smaller horns. Adult horn length was mostly independent of body mass. For adult males, the coefficient of variation of horn length (0.06) was <50% of that for body mass (0.16), suggesting that horn length presents a lower potential for selection and may be less important for male mating success than is body mass. Surprisingly, early development did not affect adult mass because of apparent compensatory growth. We found few differences in body and horn size between hunted and protected populations, suggesting the absence of strong effects of hunting on male phenotype. If horn length has a limited role in male reproductive success, hunter selectivity for males with longer horns is unlikely to lead to an artificial selective pressure on horn size. These results imply that the potential evolutionary effects of selective hunting depend on how the characteristics selected by hunters affect individual reproductive success.     

Early weaning in bighorn sheep, Ovis canadensis affects growth of males but not of females

Theories of parental investment and parent-offspring conflictassume that investment involves a cost to the parent and a benefitto the offspring, but for herbivorous mammals, behavioral andnutritional weaning are gradual processes that are difficultto define, and little is known about the consequences of individualvariation during weaning. To study the effects of late maternalcare on offspring fitness, we removed female bighorn sheep (Oviscanadensis) from a marked population in Alberta, Canada, andmonitored the survival, growth, and reproductive success oforphan and nonorphan lambs. Mothers were removed when lambswere 3.5–4.0 months, about 2–4 weeks before thesuspected time of nutritional weaning. Femaleorphans and nonorphanshad the same weight as yearlings, the same probability of producingtheir first lamb at 2 years of age, the same lifetime reproductivesuccess (lambs produced or lambs that survived to early autumn),and the same longevity. Male orphans from most cohorts weresmaller as yearlings compared to nonorphans from the same cohort.They were unable to compensate for this early weight differencein later life: at 4 years, orphan males had smaller horns andwere lighter than nonorphans. Small horn and body size likelylowered the reproductive success of orphaned males comparedto nonorphans from the same cohort. We suggest that in thissexually dimorphic species late maternal care is more importantfor males than for females. Because late maternal care had nomeasurable benefit for daughters, we suggest that parent-offspringconflict over the duration of maternal care may not exist formother-daughter pairs. or mother-son pairs it remains to beshown whether late maternal care involves a cost to the mother,but the assumption of a benefit to the son was met.     

Mass‐dependent reproductive strategies in wild bighorn ewes: a quantitative genetic approach

In the Ram Mountain bighorn sheep (Ovis canadensis) population, ewes differing by more than 30% in body mass weaned lambs with an average mass difference of only 3%. Variability in adult body mass was partly due to additive genetic effects, but inheritance of weaning mass was weak. Maternal effects could obscure genetic effects in the phenotypic expression of weaning mass, particularly if they reflected strategies of maternal expenditure that varied according to ewe mass. We performed a quantitative genetic analysis to assess genetic and environmental influences on ewe mass and on maternal expenditure. We used the mean daughters/mother regression method and Derivative Free Restricted Maximum Likelihood models to estimate heritability (h²) of ewe mass and indices of maternal expenditure. We found additive genetic effects on phenotypic variation in maternal mass, in lamb mass at weaning (absolute maternal expenditure) and in weaning mass relative to maternal mass at weaning (relative maternal expenditure). Heritability suggests that maternal expenditure has the potential to evolve. The genetic correlation of ewe mass and absolute maternal expenditure was weak, while ewe mass and relative maternal expenditure were strongly negatively correlated. These results suggest additive genetic effects on mass‐dependent reproductive strategies in bighorn ewes. Mass‐dependent reproductive strategies could affect lamb survival and phenotypic variation in adult mass. As population density increased and reproduction became costlier, small females reduced maternal expenditure more than large females. Constraints on reproductive strategy imposed by variations in resource availability are therefore likely to differ according to ewe mass. A general trend for a decrease in maternal expenditure relative to maternal size in mammals suggests that size‐dependent negative maternal effects may be common.     

Relative allocation to horn and body growth in bighorn rams varies with resource availability

Males may allocate a greater proportion of metabolic resourcesto maintenance than to the development of secondary sexual characterswhen food is scarce, to avoid compromising their probabilityof survival. We assessed the effects of resource availabilityon body mass and horn growth of bighorn rams (Ovis canadensis)at Ram Mountain, Alberta, Canada over 30 years. The number ofadult ewes in the population tripled during our study, and theaverage mass of yearling females decreased by 13%. We used theaverage mass of yearling females as an index of resource availability.Yearling female mass was negatively correlated with the bodymass of rams of all ages, but it affected horn growth only duringthe first three years of life. Yearly horn growth was affectedby a complex interaction of age, body mass, and resource availability.Among rams aged 2–4 years, the heaviest individuals hadsimilar horn growth at high and at low resource availability,but as ram mass decreased, horn growth for a given body massbecame progressively smaller with decreasing resource availability.For rams aged 5–9 years, horn growth was weakly but positivelycorrelated with body mass, and rams grew slightly more hornfor a given body mass as resource availability decreased. Whenfood is limited, young rams may direct more resources to bodygrowth than to horn growth, possibly trading long-term reproductivesuccess for short-term survival. Although horn growth of olderrams appeared to be greater at low than at high resource availability,we found no correlation between early and late growth in hornlength for the same ram, suggesting that compensatory horn growthdoes not occur in our study population. Young rams with longerhorns were more likely to be shot by sport hunters than thosewith shorter horns. Trophy hunting could select against ramswith fast-growing horns.     

Determinants of reproductive success in female Columbian ground squirrels

Summary We studied the reproductive success of female Columbian ground squirrels (Spermophilus columbianus) in southwestern Alberta for nine years. We defined reproductive success as the number of offspring surviving their first hibernation, classified as yearlings. The number of weaned juveniles explained one third of the variance in number of yearlings at emergence from their first hibernation the following spring, and much of the variance in individual reproductive success originated after weaning. Weight of adult females at emergence from hibernation was correlated with annual reproductive success. The mother's survival beyond weaning and the subsequent winter's snow accumulation had positive effects on annual reproductive success, whereas population density and summer temperature had negative effects. We found no effects on annual reproductive success of date of litter emergence, weight at emergence as a yearling, presence or absence of adult kin, distance from the natal site, location within the study area, winter temperature or summer precipitation. Age of first breeding did not affect lifetime reproductive success, which ranged from 0 to 19 yearlings produced over a lifetime. The greatest source of variation in lifetime reproductive success for females surviving to breeding age was offspring survival, followed by reproductive lifespan.     

Determinants and long‐term costs of early reproduction in males of a long‐lived polygynous mammal

In long‐lived polygynous species, male reproductive success is often monopolized by a few mature dominant individuals. Young males are generally too small to be dominant and may employ alternative tactics; however, little is known about the determinants of reproductive success for young males. Understanding the causes and consequences of variability in early reproductive success may be crucial to assess the strength of sexual selection and possible long‐term trade‐offs among life‐history traits. Selective pressures driven by fluctuating environmental conditions may depend on age class. We evaluated the determinants of reproduction in male bighorn sheep (Ovis canadensis) aged 2–4 years using 30 years of individual‐level data. These young males cannot defend estrous ewes and use alternative mating tactics. We also investigated how the age of first detected reproduction was correlated to lifetime reproductive success and longevity. We found that reproductive success of males aged 3 years was positively correlated to body mass, to the proportion of males aged 2–4 years in the competitor pool, and to the number of females available per adult male. These results suggest that reproductive success depends on both competitive ability and population age–sex structure. None of these variables, however, had significant effects on the reproductive success of males aged 2 or 4 years. Known reproduction before the age of five increased lifetime reproductive success but decreased longevity, suggesting a long‐term survival cost of early reproduction. Our analyses reveal that both individual‐level phenotypic and population‐level demographic variables influence reproductive success by young males and provide a rare assessment of fitness trade‐offs in wild polygynous males.     

Reproductive costs of sons and daughters in Rocky Mountain bighorn sheep

Differential maternal investment theory predicts that in sexuallydimorphic and polygynous species mothers should invest morein sons than in daughters. We tested the hypothesis that bighornewes that raise sons incur greater reproductive costs than ewesthat raise daughters. Although ewe mass gain during lactationand subsequent winter body mass loss were independent of lambsex, lambs born the year following the weaning of a son hadlower survival than lambs born after a daughter. The effectsof lamb sex on subsequent reproductive success of ewes becamemore evident at high population density. Lamb sex did not affectmaternal survival. Population density, weather, and ewe agedid not alter the relationship between lamb sex and subsequentreproductive success of the ewe. The year after weaning a son,ewes were more likely to have a daughter than a son, while ewesthat had previously weaned a daughter had similar numbers ofsons and daughters. Our results show that for bighorn sheepewes, sons have a greater life-history cost than daughters,suggesting a differential maternal investment in the sexes.     

Determinants of age at first reproduction and lifetime breeding success revealed by full paternity assignment in a male ungulate

Age at first reproduction is an important determinant of individual variation in reproductive success in ungulates, but few studies have examined its relationship with later fitness‐related traits in males. We used a long‐term individual based study of a harvested moose population to quantify the individual reproductive performance and survival of males, as well as to examine the determinants of age at first reproduction and consequences of age at first reproduction on lifetime breeding success. The probability that a male successfully reproduced at the age of two was negatively related to the mean age of adult males in the population, but the relationship weakened with increasing population size. Large antlers and large body mass relative to other males in the population increased the number of calves sired at their first successful mating season. In addition, those that successfully reproduced as two year‐olds were more likely to sire calves the next year, making them more productive at a given age compared to those that first reproduced at the age of three or older. We emphasize the importance for males to start reproducing as soon as possible in a harvested population to gain lifetime fitness benefits, as surviving the hunt is a major determinant of reproductive success in this population. We found no costs of early reproduction in males, hence leading to high individual heterogeneity in male reproductive performance. The apparent lack of reproductive costs could partly be explained by the age distribution in the population, individual variation in early‐life body mass and antler size, and differences in probabilities of being hunted of successful and unsuccessful males.     

Sex- and age-specific survival of the highly dimorphic Alpine ibex: evidence for a conservative life-history tactic

1. Age-specific survival of 215 males and 117 females of the highly sexually dimorphic Alpine ibex Capra ibex (L.) was assessed from a 21-year capture-mark-recapture (CMR) programme (1983-2004). The study covered two contrasted periods of population performance (high performance from 1983 to 1997 vs. low performance from 1998 onwards). 2. Based on current life-history theories for sexually dimorphic species, we expected that survival should decrease with age in both sexes, female survival should be buffered against environmental variations, male survival should decrease during the low performance period, and adult survival should be lower in males than females during the low performance period. 3. Survival of both sexes was strongly affected by age, with the four age classes (yearling, prime-aged adults of 2-8 years of age, old adults of 8-13 years of age, and senescent adults from 13 years of age onwards) generally reported for large herbivores. 4. Survival of females at all ages, and of yearling and prime-aged males, was buffered against environmental variations and was the same during periods of high and low population performance. The survival of old males decreased in years of low population performance. 5. All marked yearlings (32 females, 56 males) survived to age 2. Survival of prime-aged females (0.996 +/- 0.011) was higher than for other large herbivores, but similarly to other large herbivore species, it declined slowly and regularly with increasing age afterwards. Male survival was 5-15% higher each year than that of males of other large herbivores. Males enjoyed very high survival when prime-aged (0.981 +/- 0.009) and as old adults (high-performance period: 0.965 +/- 0.028, low-performance period: 0.847 +/- 0.032). 6. The very high survival of males, coupled with their prolonged mass gain, suggests a highly conservative reproductive tactic. Male ibex differ from similar-sized herbivores by showing a nearly indeterminate growth in horn size and body mass. By surviving to an advanced age, males may enjoy high reproductive success because of their large size.     

A selective trade-off for territoriality and non-territoriality in the polymorphic damselfly Mnais costalis

Males of the damselfly Mnais costalis occur as territorial orange-winged 'fighter' males or non-territorial clear-winged 'sneaker' males. Their morph life histories differ considerably but the estimated lifetime reproductive success is the same for the two morphs. In this study we compared the developmental and reproductive costs associated with the two morphs. Orange-winged male and female reproductive costs resulted in a decline in adult fat reserves with increasing age. In contrast, the fat reserves of clear-winged males remained constant with adult age. Body size was positively correlated with mating success in orange-winged males, but had no influence on the mating success of clear-winged males. The orange-winged male flight muscle ratios (FMRs) were significantly higher than the clear-winged male and female FMRs. However, there was no difference in the size-corrected fat reserves of the two morphs; both had higher fat reserves than females. The gain in mass between eclosion and reproduction in orange-winged males and females was almost double the mass gained by clear-winged males, suggesting that clear-winged male development is less costly. An experiment in which pre-reproductive levels of nutrition were manipulated confirmed this.     

Father–offspring phenotypic correlations suggest intralocus sexual conflict for a fitness-linked trait in a wild sexually dimorphic mammal

In sexually dimorphic and polygynous mammals, sexual selection often favours large males with well-developed weaponry, as these secondary sexual characters confer advantages in intrasexual competition and are often preferred by females. Little is known, however, about the effects of sexually selected paternal traits on offspring phenotype in wild mammals, especially when considering that shared phenotypic traits and selection can also differ greatly between genders. Here, we conducted molecular parentage analyses in a long-term study population of mountain goats (Oreamnos americanus), an ungulate exhibiting high sexual dimorphism in mass, to first assess the determinants of yearly reproductive success (YRS) in males. We then examined the effects of paternal characteristics on offspring mass at 1 year of age. Paternity was highly skewed, with 9 per cent of 57 males siring 51 per cent of 96 offspring assigned over 12 years. Male YRS increased with age until apparent reproductive senescence at 9 years, but mass was a stronger determinant of siring success than age, horn length or social rank. Mass of sons increased with paternal mass, but the mass of daughters was negatively related to that of their father, a finding consistent with recent theory on intralocus sexual conflict. Because early differences in mass persisted to early adulthood, sex-specific effects of paternal mass can have important fitness consequences, as adult mass is positively linked with reproduction in both sexes. Divergent father–offspring phenotypic correlations may partly explain the maintenance of sexual dimorphism in mountain goats and the large variance observed for this homologous trait within each gender in polygynous mammals.     

Scramble competition polygyny in the milkweed leaf beetle: combat, mobility, and the importance of being there

I examined the scramble competition mating system of the milkweedleaf beetle, Labidomera clivicollis(Coleoptera: Chrysomelidae),using data on lifetime mating success of two breeding cohortsin Austin, Texas, USA (30° N, 97° W) and one in Bridgeport,New York, USA (43° N, 76° W). Data from six daily censuseswere combined with focal animal samples to examine the importanceof combat between males and to evaluate the extent to whichthe characters elytron length, vagility, and emergence datewere correlated with male mating success. In both populations,emergence date was the only consistent determinant of the numberof mates a male obtained during his lifetime. Males emergingearlier in the season had higher mating success because theywere active for more of the breeding season. Male body size(elytron length) was not correlated with mating efficiency orlifetime mating success in either population. Males formed prolongedmating associations with females that lasted for up to 2.5 daysand paired males in the New York population had a strong advantageover interlopers attempting to mate with the female. Only 10%of takeover attempts were successful. Selection favored vagilemales in New York, where the sex ratio was male biased, butnot in Texas, where it was female biased. Males that were morevagile had higher mating efficiencies in New York but did nothave higher lifetime mating success because of a trade-off betweenmating efficiency and survival. Behavioral data demonstratedthat lone males spent more time walking and less time feedingthan lone females; mating males were unable to feed at all.The survival disadvantage that comes with increased vagilitymay be due to loss of time spent feeding. The characters examinedsuggest that the most important thing a male L. clivicolliscan do to increase his lifetime mating success is to be presentfor as much of the breeding season as possible.     

Correlates of reproductive success within alternative mating tactics of the common shrew

Male common shrews (Sorex araneus) adopt two discrete matingtactics. The most successful males, in terms of number of offspringfathered, are those that establish large overlapping home rangesin areas of high female density early in the breeding season.An alternative, less successful mating tactic is to travel longdistances in search of mating opportunities. This study is aninvestigation of correlates of reproductive success for malesadopting these different mating tactics. Reproductive successunder natural conditions was assessed using DNA fingerprinting,and survival of offspring was monitored in the field. The numberof offspring fathered by males with overlapping home rangeswas positively correlated with the number of female ranges overlappedduring the breeding season and with testes mass. The numberof offspring fathered by males that made long-distance movementswas positively correlated with their epididymal sperm counts.It is argued that competitively superior (overlapping) malesachieve high reproductive success by competing to maximize thenumber of females inseminated, whereas those adopting an alternativemating tactic instead compete largely via sperm competition,aiming to maximize insemination success with any particularfemale. There was no significant difference in the fluctuatingasymmetry (FA) of males adopting different mating tactics (FAwas measured as the difference in length of the paired lateralscent glands). Fluctuating asymmetry was not related to thenumber of offspring fathered by males adopting either matingtactic, but was significantly correlated with the proportionof male offspring fathered that survived to sexual maturity.Although apparently not correlated with mating success in thisspecies, FA may reflect some aspect of genetic quality thataffects offspring survival. [Behav Ecol 7: 334–340 (1996)]     

Live fast, die young: trade-offs between fitness components and sexually antagonistic selection on weaponry in Soay sheep

Males are predicted to compete for reproductive opportunities, with sexual selection driving the evolution of large body size and weaponry through the advantage they confer for access to females. Few studies have explored potential trade-offs of investment in secondary sexual traits between different components of fitness or tested for sexually antagonistic selection pressures. These factors may provide explanations for observed polymorphisms in both form and quality of secondary sexual traits. We report here an analysis of selection on horn phenotype in a feral population of Soay sheep (Ovis aries) on the island of Hirta, St. Kilda, Scotland. Soay sheep display a phenotypic polymorphism for horn type with males growing either normal or reduced (scurred) horns, and females growing either normal, scurred, or no (polled) horns; further variation in size exists within horn morphs. We show that horn phenotype and the size of the trait displayed is subject to different selection pressures in males and females, generating sexually antagonistic selection. Furthermore, there was evidence of a trade-off between breeding success and longevity in normal-horned males, with both the normal horn type and larger horn size being associated with greater annual breeding success but reduced longevity. Therefore, selection through lifetime breeding success was not found to act upon horn phenotype in males. In females, a negative association of annual breeding success within the normal-horned phenotype did not result in a significant difference in lifetime fitness when compared to scurred individuals, as no significant difference in longevity was found. However, increased horn size within this group was negatively associated with breeding success and longevity. Females without horns (polled) suffered reduced longevity and thus reduced lifetime breeding success relative the other horn morphs. Our results therefore suggest that trade-offs between different components of fitness and antagonistic selection between the sexes may maintain genetic variation for secondary sexual traits within a population.     

密点麻蜥的两性异形和雌性繁殖

蜥蜴繁殖成功率与其形态特征有密切的关系。作者在内蒙古乌拉特后旗采集密点麻蜥(Eremias multio-cellata) ,定量研究该种形态特征的两性异形和雌体繁殖特征,检验与成体形态特征相关的两性繁殖成功率差异是否能促进两性异形的进化。密点麻蜥成体个体大小无显著的两性差异,但头部大小两性差异显著;雄性个体的头长和头宽均大于体长相同的雌性成体。繁殖雌体于五、六月份排卵;在实验室条件下,雌体在六月下旬至七月下旬之间产仔。该种雌体年产单窝仔,每窝2 -4仔。窝仔重与雌体体长呈正相关,但雌体体长仅能解释很少一部分(约19 %)窝仔重的变异。窝仔数和幼仔重均与雌体体长无关。幼仔重与相对生育力(相对于雌体体长的窝仔数)呈显著的负相关,表明该种蜥蜴存在后代数量-大小之间的权衡。密点麻蜥雄体和雌体向较大体型方向进化的选择压力均相对较弱,与成体头部大小相关的两性繁殖成功率的差异是导致该种蜥蜴头部大小两性异形进化的主要原因[动物学报52 (2) : 250 -255 , 2006]。     

Maternal expenditure in the polygynous and monomorphic guanaco: suckling behavior, reproductive effort, yearly variation, and influence on juvenile survival

We investigated patterns of maternal expenditure and its influenceon juvenile survival in the polygynous monomorphic guanaco (Lama guanicoe)in southern Chile from 1990 to 1994. Birth weight and growth rate(until age 1) of males and females were similar. Suckling ratesof males and females were not significantly different, althoughmothers of males rejected suckling attempts more often thanmothers of females during fall and winter. Mothers with sonsterminated suckling bouts in equal proportion as did motherswith daughters. Our estimated level of reproductive effort for guanacosfalls within the range of species exhibiting no sex-biased maternal expenditureon offspring. Mean yearly birth weight was negatively correlated withpopulation density. Mean suckling time throughout the year differedamong cohorts, as did the mean number of suckling attempts andrejected suckling attempts per hour throughout the year. Juvenilesurvival was estimated until age 1. Of the model with five covariatesincluding juvenile sex, birth weight, adult female aggressiontoward taggers, mean suckling time, and population density,only mean suckling time and population density were significantly relatedto survival. The risk ratio for mean suckling time indicatesthat the risk of mortality increases as suckling time increases,whereas the risk ratio for population density indicates thatthe risk of mortality decreases as population density increases.Under some conditions increasing population density may be correlatedwith lower offspring birth weight, yet enhanced juvenile survival.This effect on survival was possibly associated with the numberof predators on the study area from year to year.     

Sexual selection in flour beetles: the relationship between sperm precedence and male olfactory attractiveness

Sperm precedence, defined as nonrandom differential fertilizationsuccess among mating males, is an important postmating componentof sexual selection. This study examined the relationship betweenpremating and postmating components of sexual selection in malesof the flour beetle (Tribolium castanewn). Male olfactory attractivenessto females was positively correlated with a male's subsequentfertilization success: more attractive males achieved highersecond-male sperm precedence when allowed to mate with previouslyinseminated females. Attractive males may achieve compoundedgains in their reproductive success through enhanced matingopportunities as well as through greater fertilization success.Thus, the relationship between these reproductive fitness componentsmay augment differences in reproductive success among males.Female fecundity, estimated as the number of adult progeny produced,increased significantly with multiple malings. This result supportsincreased female reproductive success as a direct benefit ofmultiple mating in T. caslaneum and suggests that progeny productionis partially limited by sperm availability. Total progeny productionby doubly mated females remained constant at all levels of second-malesperm precedence. However, higher sperm precedence was associatedwith a decline in firstmale progeny and a concomitant increasein second-male progeny. This pattern of progeny production suggeststhat more attractive males may achieve higher fertilizationsuccess through a combination of displacement of previouslystored sperm, transfer of greater sperm quantities, or females'preferential use of sperm of attractive males for fertilizations.     

Life history of the moose Alces alces: relationship between growth and reproduction

We studied the relationship between increase in body weight and reproductive performance in different populations of Norwegian moose to evaluate costs associated with early onset of reproduction, viz. whether early onset of reproduction was correlated with low adult body weight or reduced adult fecundity. The mean carcass weight of non-ovulating yearlings was significantly lower than for ovulating yearlings. However, those 2.5 yr old females that conceived as yearlings were lighter than non-reproducing females of the same age. Thus, to begin to reproduce as a yearling was assumed to be expensive because it reduces the possibilities for further growth. The cost associated with reproduction was further illustrated by the fact that the difference in mean carcass weight from age 2.5 to 3.5 yr of females that produced calves in both years, was less for the females from regions with lowest mean yearling weights, i.e. regions with probably the lowest resource availability. In populations with high mean yearling carcass weights, the proportion of cows with calf and the number of calves per pregnant female in the early reproductive phase (2.5 or 3.5 yr old) were higher than in populations in which the mean yearling weights were low. There was a negative correlation between growth rate in the population after 1.5 yr of age and the mean yearling carcass weight. Thus, low yearling weight was associated with a prolonged period of growth and thereby a reduced reproductive output during the first year of the female's life. For old females (≥ 9.5 yr old) the number of calves produced per pregnant female was highest in populations where yearling carcass weights were highest. Furthermore, mean yearling weight and the mean adult female weight were positively correlated in those regions. This relationship suggests that within this species early onset of reproduction is not related to retarded reproduction or lower weight later in life. We suggest that the moose has been selected for an early onset of reproduction.