Sex differences in mass loss rate and growth efficiency in Antarctic fur seal (Arctocephalus gazella) pups at Macquarie Island

We investigated the relationship between the mass gained by a pup during a period of maternal attendance (as an index of milk intake) and the duration of the preceding foraging trip in relation to the mass-specific rate of mass loss during fasting periods and the growth rate of Antarctic fur seal (Arctocephalus gazella) pups at Macquarie Island. We found that (1) serially weighed male pups grew significantly faster than females pups and that (2) fasting female pups lost mass at a significantly higher rate (2.55% day⁻¹) than male pups (2.12% day⁻¹) of the same mass; (3) during periods of maternal attendance, there were no intersexual differences in the amount of mass gained by pups of the same size, hence (4) female pups required a higher daily mass gain to grow at the same rate as male pups. Our results show that intersexual differences in growth rate may be accounted for by intersexual differences in mass-specific rate of mass loss, because females lost 0.42% more of their total mass per day (i.e. 4.2 g kg⁻¹ day⁻¹) compared with male pups of the same body mass. Despite intersexual differences in growth rates, our results indicate equality of maternal expenditure between the sexes. Intersexual differences in the rate of mass loss may be due to differences in the metabolic rate, activity level and/or body composition of male and female pups. Received: 19 August 1998 / Received in revised form: 19 April 1999 / Accepted: 19 April 1999     

Intraspecific allometry of neonatal size in the antarctic fur seal (Arctocephalus galapagoensis)

Summary Intraspecific allometry of pup mass as a function of maternal mass was analysed in Antarctic fur seals (Arctocephalus gazella), a highly dimorphic species. The allometric exponent was 0.6, much lower than in interspecific comparisons. Slopes were the same for mother-son pairs as for mother-daughter pairs, but adjusted means were significantly higher for the former, indicating higher reproductive effort of mothers of sons. The correlation of maternal mass with pup mass explained 59% of the variance in female pup mas but only 18% of that in male pup mass. Females appeared to produce sons which were as big as possible. Relative pup mass decreased with increasing maternal mass indicating a lower reproductive effort of bigger and presumably older mothers, contrary to expectations from life history theory. Sex ratio of pups showed no relation to maternal mass.     

Maternal investment and sex-allocation in the Galapagos fur seal,Arctocephalus galapagoensis

Summary Maternal investment and sex-allocation were measured in a large, sexually dimorphic mammal, the Galapagos fur seal (Arctocephalus galapagoensis). The sex ratio at birth was 1.06. Males were always heavier than females and, at least initially, grew faster. Growth was variable from year to year suggesting energetic constraints on maternal investment. Sucking time conrrelated with milk intake. Mothers suckled yearling and 2-year-old sons more than daughters of the same age. Age at weaning appeared to be the same in both sexes or even slightly greater in males. No sex differences was found in mortality prior to weaning or in post-weaning dispersal. Birth rates of females with yearlings or 2-year-olds were significantly lower than those of females with no dependent young. Mothers invested more in sons than in daughters until weaning. It is unlikely that higher post-weaning investment in daughters balances the higher pre-weaning investment in sons. Data on sex ratio at birth, different growth rates, and weaning age of the sexes are typical of otariid seals as a group. The results of this study fit Maynard Smith's (1980) model of the evolution of sex allocation better than Fisher's (1930).     

An energy–distance trade-off in a central-place forager,the Antarctic fur seal (<Emphasis Type="Italic">Arctocephalus gazella</Emphasis>)

We tested the prediction that lactating fur seals (Arctocephalus gazella) at South Georgia will take prey of greater energy density with increasing distance of foraging from the colony. The study investigated the differences in diet of fur seals foraging within two regions, one near the breeding colony and the other at greater distance. Diet varied significantly in relation to foraging location. Dietary items of low quality were eaten in both regions but more food items with a high-energy content appeared in the diet of seals travelling to distant oceanic waters. We conclude that there is likely to be a trade-off between energy gain and distance travelled which enables female fur seals to maintain a relatively constant rate of energy delivery to their offspring irrespective of the distance travelled to find food.     

Female reproductive qualities affect male painted turtle (Chrysemys picta marginata) reproductive success

Male reproductive success (RS) in polygamous species with minimal social systems is often determined by the number of mates. However, because male RS is translated through females, the number of offspring sired can also be influenced by female qualities. Empirically sufficient data to document how tradeoffs between mate number and quality influence male RS are seldom available for long-lived, iteroparous species. We combined long-term life history data (1983–2006) on the E. S. George Reserve (ESGR, MI, USA) with parentage data from 155 clutches of 59 female painted turtles (Chrysemys picta marginata) of varying reproductive frequencies (2003–2006) to determine the relative contribution of female numbers and qualities on male RS. One previously documented trait of female painted turtles that can have substantial influences on male RS is repeat paternity through the use of stored sperm to fertilize over 95 % of within-year clutches. In addition, our study found that second-clutch producing female painted turtles on the ESGR have higher among-year reproductive frequencies than do first-clutch only females. Multiple paternity was detected in 14.1 % of clutches (min-max?=?6.1–30.0 % annually), and the number of mates of both sexes was low annually (males 1.0; females 1.2) and over 4 years (males 1.1; females 1.7). Among successful males, RS varied substantially (1–32 offspring) and was strongly influenced by the combination of female reproductive frequency and repeat paternity (>38 % among years), but not mate number. Low mate number for both sexes was unexpected in a species without complex mating behaviors or parental care.     

Mitochondrial DNA sequence data of the Cape fur seal (Arctocephalus pusillus pusillus) suggest that population numbers may be affected by climatic shifts

The Cape fur seal, Arctocephalus pusillus pusillus is distributed along the southern African coastline from northern Namibia to the south-east coast of South Africa. The species has been impacted by sealing operations since the 1600s, and historical records suggest that the taxon experienced a bottleneck prior to the 20th century. Mitochondrial DNA control region sequences were generated for 106 individuals belonging to six breeding colonies. Haplotype diversity was found to be high (0.975±0.014) whereas levels of nucleotide diversity were much lower compared to other seal species (0.011±0.006). An analysis of molecular variance indicated that the largest percentage of haplotype diversity is distributed within colonies rather than among them. This could be attributed to either extensive gene flow among colonies, a lack of substantial female site philopatry, or incomplete lineage sorting of haplotypes. Mismatch distribution and Fu’s F _S test indicated that the population has experienced a historical population expansion probably between c. 37,000–18,000 YBP and this date coincides very well with the height of the last glacial maximum when food resources were abundant in the South Atlantic. These results also suggest that the recent sealing-induced bottleneck did not have a profound influence on the haplotype diversity and a historical bottleneck prior to a demographic expansion may have been severe enough to reduce nucleotide diversity substantially.     

Individual and environmental determinants of reproductive success in male tree swallow (Tachycineta bicolor)

Evaluating the contribution of individual and environmental determinants of reproductive success is essential to improve our understanding of sexual selection. In socially monogamous bird species with high rates of extrapair paternity, traits or environmental contexts affecting the number of within-pair young (WPY) produced by males can differ from those affecting the number of extrapair young fathered (EPY). Here, we use a 4-year dataset collected in contrasted environments to assess the factors affecting male reproductive success in tree swallows (Tachycineta bicolor), a species showing high levels of extrapair paternity. Our analyses revealed that the number of WPY was higher under better environmental conditions, while the number of EPY was mainly related to male characteristics. Males nesting in more intensive agricultural areas had fewer WPY produced and a lower reproductive success. Also, males breeding earlier in the season had more WPY. The presence of parasites reduced males’ reproductive success, mainly by reducing the number of EPY. The influence of male phenotype varied according to population density: Tarsus length variation had a greater effect on reproductive success at low population density than at high density, while wing length was also positively related to the number of EPY, more so at high than at low density. Altogether, our results suggest a complex interplay between individual and environmental determinants of reproductive success and imply that sexual selection dynamics varies depending on environmental contexts.     

Sex-specific patterns of lifetime reproductive success in single and repeat breeding steelhead trout (Oncorhynchus mykiss)

All other things being equal, the lifetime reproductive success (LRS) of iteroparous and semelparous individuals should scale with the number of breeding seasons. Deviations from this relationship may occur for many reasons, including age- or size-related fecundity or life history trade-offs, which may differ between sexes. We used 19 brood years of DNA parentage analysis in a small (N = 4–143 year⁻¹) wild, unexploited population of steelhead trout (Oncorhynchus mykiss) to compare the LRS of individuals that spawned only once [“one time spawners” (OTS), N = 355 male, 371 female] to those spawning twice [“repeat spawners” (RPS), N = 13 male, 49 female]. Female RPS had nearly twice the LRS of female OTS (1.17 offspring per female vs 0.91 offspring per female), whereas male RPS had nearly three times the LRS of male OTS (1.54 offspring per male vs 0.57 offspring per male). Female RPS produced slightly more adult offspring during their second breeding season than their first (0.78 vs 0.82 offspring per female); however, male RPS produced all of their adult offspring in their second breeding season (0 vs 1.54 offspring per male). The additional growth in body size of males between breeding seasons may give them an advantage in their second breeding season, but the lack of offspring produced in their first season suggests a trade-off between survival and future reproduction that was not expressed in females.     

Factors affecting male and female reproductive success in a chipmunk (Tamias sibiricus) with a scramble competition mating system

While sexual selection in mammals with female defense systems has historically received considerable attention, much less is known about the factors affecting reproductive success in mammalian species with scramble competition mating systems. Using mark–recapture techniques and DNA microsatellite loci to assign parentage, we examined the mating system and factors affecting the variation of the annual reproductive success in an introduced population of the Siberian chipmunk, Tamias sibiricus, a small, monomorphic, solitary squirrel. Our analyses showed that breeding females were spatially dispersed, setting the stage for a scramble competition mating system. Male reproductive success was positively associated with the size of the home range. The strength of sexual selection on this behavioral trait was very strong, equaling previous estimates for morphological traits in female defense mating systems. These findings suggest that a behavioral trait, space use, strongly influences the annual reproductive success in males, which is consistent with expectations in a scramble competition mating system. In both sexes, reproductive success was influenced by habitat, with twice as many juveniles produced in semi-open than in closed habitat, possibly due to differences in food availability between the two habitats.     

Sigma virus and male reproductive success in Drosophila melanogaster

The risk of disease transmission can affect female mating rate, and thus sexual conflict. Furthermore, the interests of a sexually transmitted organism may align or diverge with those of either sex, potentially making the disease agent a third participant in the sexual arms race. In Drosophila melanogaster, where sexual conflict over female mating rate is well established, we investigated how a common, non-lethal virus (sigma virus) might affect this conflict. We gave uninfected females the opportunity to copulate twice in no-choice trials: either with two uninfected males, or with one male infected with sigma virus followed by an uninfected male. We assessed whether females respond behaviorally to male infection, determined whether male infection affects either female or male reproductive success, and measured offspring infection rates. Male infection status did not influence time to copulation, or time to re-mating. However, male infection did affect male reproductive success: first males sired a significantly greater proportion of offspring, as well as more total offspring, when they were infected with sigma virus. Thus viral infection may provide males an advantage in sperm competition, or, possibly, females may preferentially use infected sperm. We found no clear costs of infection in terms of offspring survival. Viral reproductive success (the number of infected offspring) was strongly correlated with male reproductive success. Further studies are needed to demonstrate whether virus-induced changes in reproductive success affect male and female lifetime fitness, and whether virus-induced changes are under male, female, or viral control.     

Determinants of male reproductive success in American black bears

We determined annual male reproductive success in black bears (Ursus americanus) using DNA and field data from two populations in New Mexico. We identified the likely father for 78 of 120 genotyped cubs born during 1994–2000 and calculated reproductive success for 102 males known or presumed present within designated mating regions. Age was a strong determinant of reproductive success. The observed peak in reproductive success occurred at roughly the same intermediate age (10 years) as milder peaks in body size characteristics (11–12 years) and frequency of bear-inflicted wounds (13 years), suggesting body size and fighting might be important for mating. Success was negatively associated with the distance between home range centers of males and mothers. Success of young males (<7 years old) was also negatively associated with mature male (≥7 years old) density, and increasing density shifted the peak age of reproduction higher. The dispersed distribution of females likely limited the capacity of large mature males to dominate reproduction; therefore, success was determined by a complex set of variables.     

Importance of monogamous male birds in determining reproductive success

Summary Several recent studies have questioned whether the presence and activities of monogamous male birds are of value to their young during the nesting attempt. We addressed this issue in house wrens (Troglodytes aedon) by removing males early in the nestling stage. In one of four periods survivorship among experimental nestlings was 63% lower than among control nestlings. In the other periods survivorship was about equal in experimental and control nests indicating that in this species the male's activities may only benefit his nestlings during unfavorable periods. We summarize the results of 15 other male removal studies, concluding that when males appear to be helping, their absence usually results in decreased survival of young whereas when the males render little apparent care their absence does not decrease survival of the young. A possible exception to this generalization, illustrated by our study and others, is that during the nestling stage of many altricial species, the female appears able to raise the young about as well without the male as with him, except in poor periods. A male caring for his nestlings during good periods may benefit by maintaining his mate's health for the next nesting attempt or by insuring her willingness to mate with him again. Alternatively, it may cost males so little to help during good periods, and benefit them so much during poor periods, that there has been little selective pressure for them to assess conditions and vary their level of care accordingly. A series of more specific predictions about which males are likely to be of greatest, and least, value to their nestlings is presented. Male care after young leave the nest has not been well studied, but is probably substantial in many species, and warrants attention in future studies.     

Fieldfare (Turdus pilaris) breeding success in relation to colony size,nest position and association with Merlins (Falco columbarius)

Summary Clutch size, nestling production and breeding success were studied in colonial Fieldfares (Turdus pilaris) in a subalpine birch forest during ten breeding seasons. Reproductive success was highest for central pairs in large colonies; such pairs benefited most from communal defence against nest predators. Fieldfares and Merlins (Falco columbarius) usually bred in association. Fieldfares breeding away from Merlins had lower breeding success than pairs associated with Merlins, which also benefited by reduced nest predation. Fieldfares apparently chose to nest near Merlins, which had already laid eggs when the thrushes started nest-building.     

Correlates of male reproductive success in the browncheek blenny,Acanthemblemaria crockeri (Blennioidea: Chaenopsidae)

Summary Acanthemblemaria crockeri exhibits a resource defense polygyny mating system with male parental care. Females preferred to mate with the larger of two males in laboratory experiments, and male size was positively correlated with the number of eggs defended by males at two sites in the Gulf of California, Baja California, Mexico. Females appeared to avoid mating with males defending heavily-fouled shelters. The role of other factors including the intensity of male courtship coloration and displays in determining male reproductive success was studied in Bahia San Carlos, Sonora, Mexico by providing glass vials as shelters. This provided a non-destructive technique for counting of eggs defended by males and allowed repeated assessment of the reproductive success of individual males. Males varied greatly in their duration of residency of shelters and in their color score and intensity of courtship displays. The number of eggs received by males was positively correlated with their duration of residency in shelters, and females appeared to avoid mating with new residents. Mated males deserted shelters less frequently and were more likely to receive future matings than were unmated males. The mean color score of males was unrelated to their reproductive success, while the intensity of male courtship displays was negatively correlated with the number of eggs received. This may have resulted because female mate choice is based on multiple criteria, including some that more accurately reflect the quality of parental care afforded by males.     

The reproductive behavior and energetics of male gray seals (Halichoerus grypus) breeding on a land-fast ice substrate

The reproductive behavior of male gray seals (Halichoerus grypus) breeding on land-fast ice at Amet Island, Nova Scotia, was studied. Data on energy expenditure (rate of mass loss over time) were collected. The average time budget of males at Amet Island was comparable to that of land-breeding males. The behavior of males showed seasonal changes, with a decrease in the proportion of time spent in the water and an increase in agonistic behavior during the peak mating period. The estimated amount of body mass lost over the season ranged between 25.6 and 77.1 kg, and the estimated percent of initial body mass lost ranged between 7.7 and 26.5% (n=10). The maximum number of observed copulations for an individual male was nine. Only 15 out of 42 males observed during 1992 and 1993 were seen copulating. The number of observed copulations per male was strongly correlated with success in remaining close to, or attending, females (r=0.91, P<0.001, n=42). The mean duration of attendance was 4.5 ± 5.54 days (n=42). Large size was not an important factor in determining attendance success, but reproductive effort (the estimated proportion of body mass lost over the season) and success in agonistic interactions with other males were both correlated with male success.     

The hunting strategy of white sharks (Carcharodon carcharias) near a seal colony

The degree to which white sharks, Carcharodon carcharias, are social while hunting is unclear. Our aim was to describe the behavior and interactions among white sharks hunting seals near a seal colony. We attached ultrasonic beacons to five adult white sharks, 4.5–5.2?m long, and recorded their movements and behavior toward each other over a 15-day period in October 1997 at Año Nuevo Island, California. This site is home to colonies of four species of seals and sea lions. Two additional sharks, females 5.5 and 4.7?m in length, were later tracked intensively during periods of 12 and 3?days during October 1998 and November 1999, respectively. We recorded stomach temperature (indicative of feeding on warm-bodied seals) and swimming depths from the 5.5-m female, swimming speed and depth from the 4.7-m female. We monitored the movements and behavior of these sharks using an array of sonobuoys moored near the island; the receptive field measured 1?km². Our principal findings were: (1) the sharks spent a mean time of 39.5% of each day patrolling within the receptive field; (2) no shark ever moved far out of it; (3) the sharks spent an equal amount of time and activity in the receptive field at all times of the day, daytime, twilight, and nighttime; (4) movements with respect to the island rookery were most often back and forth parallel to the shoreline, (5) tracks of three sharks, tagged at the same time and place, overlapped more often than those of the other two sharks; and (6) some sharks patrolled certain areas in the field preferentially, but there was no conclusive evidence that they defended these areas as territories. Feeding appeared to be infrequent: only two likely feeding bouts occurred during a cumulative 78-day/shark period that individuals were monitored at Año Nuevo Island. The behavior and movements of the sharks were consistent with a hunting strategy, in which individuals search for prey independently but, at the same time, remain close enough to each other to “sense” and exploit a kill by any one of them by joining in on the kill to feed.     

Components of variance in male lifetime copulatory and reproductive success in a seed bug

Summary Males and females of the seed bug, Neacoryphus bicrucis Say, were individually numbered in the field in southeastern Georgia (USA) and census taken daily for 6 weeks. Variation in male mating efficiency (ME = no. copulations/no. sightings) exceeded that in females and was significantly greater than that generated by a null model. Lifetime copulatory success, estimated as the product of ME and longevity, ranged from 0 to 41, with ME accounting for over 88% of the variation. Lifetime reproductive success (LRS), estimated as the product of ME, longevity, and clutch size, ranged from 0 to 898. Among males copulating at least once, ME accounted for over 40% of the variation in LRS, while longevity alone or in combination with clutch size accounted for 21% and 46%, respectively, of the variation in LRS. Failure of some males (26%) to copulate contributed 38% of the total variation in LRS. Thus, among males surviving to adulthood, sexual selection pressure arising from variation in ME is approximately as strong a force for phenotypic change as is natural selection pressure arising from variation in longevity.     

Female habitat choice as a determinant of the reproductive success of the territorial male marine iguana (Amblyrhynchus cristatus)

Summary The reproductive behaviour of marine iguanas (Amblyrhynchus cristatus) was studied to answer the following questions: Which physical factors determine the number of females found on a male's territory? Do males select territories according to preferences for physical conditions or according to the number of females there? What causes the strong correlation between the number of females on a territory and the number of copulations achieved there? Females preferred as their resting places territories with the following characteristics, in order of importance: High number of shady/shelter places, low temperature of rocks at noon, proximity to the sea, sun-basking places in the afternoon. Other factors measured (probability of being wetted by spray, sun-basking places in the morning and at noon) showed no significant correlation with the number of females. Males, however, preferred territories further from the sea (within about 7 m), with high rock temperatures and a high number of basking places in the morning (in order of importance). Males primarily followed their site preferences and not female density in their choice of a territory. The site choice of females determined the number of copulations achieved on a territory, as females copulated preferentially on their resting places. A shift of females from one territory to another within one colony between two succeeding years was correlated with the change in relative weight of territory owners. This result was interpreted as an indication of female choice. Site preferences of males and females are discussed in relation to the animals' physiological requirements. Possible influences of ecological factors on the evolution of this mating system are considered.     

Variance in male and female reproductive success in a harem-polygynous mammal,the black-tailed prairie dog (Sciuridae: Cynomys ludovicianus)

Summary Black-tailed prairie dogs are colonial rodents that live in contiguous social groups called coteries. A typical coterie contains one adult (2 years old) male, three or four adult females, and several yearlings and juveniles of both sexes. A large coterie sometimes contains two or more adult males. Using detailed behavioral observations on 164 females (of which 160 copulated) and data from four polymorphic loci from parents and offspring of 121 litters, we examined the black-tail mating system.Most females (101/164=62%) copulated with a single adult male, and only 3 of the 102 litters with 2 offspring (3%) showed unequivocal evidence of multiple paternity. Adult males usually copulated with several different adult females.In one-male coteries, females usually copulated exclusively with the resident adult male (RAM) (82/112=73%); this trend was confirmed by electrophoresis of blood smaples. In multimale coteries, each female frequently copulated with at least two different RAMs (28/52=54%); in 4 of 5 multimale coteries (80%) which produced two or more litters whose paternities could be unequivocally resolved by electrophoresis, two different RAMs each sired at least one litter.Of the 164 females, 30 (18%) copulated with both the RAM (or one of the RAMs, in multimale coteries) and an extracoterie adult male, but only 3 (2%) copulated exclusively with an extracoterie adult male. Electrophoresis showed that 9 of 121 litters (7%) were sired by an extracoterie adult male.Intersexual comparisons of annual reproductive success and lifetime reproductive success both indicate that black-tails are polygynous (i.e., that variance in reproductive success is greater for males than for females).