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.     

Environmental predictability and remating in European blackbirds

We studied mate and site fidelity between years in an Englishpopulation of blackbirds (Turdus merula) from 1985 to 1991.Divorce was observed in 32% of 183 cases where mates had survivedfrom the previous breeding season. After divorces, females tendedto nest farther from their original site than males did, butthere was no sex difference in distance moved after the deathof a mate. The mean annual fledgling production of 60 m x 60m breeding sites was statistically repeatable among years, evenafter accounting for possible differences in reproductive performanceamong site-tenacious individuals. Divorce rates were greatestin low-quality sites. We found no annual fitness cost associatedwith mating with an unfamiliar bird. However, breeding in anew site cost on average one fledgling per year.     

Seasonal changes in condition, reproduction and fecundity in the wild European rabbit (Oryctolagus cuniculus)

A total of 1248 rabbits ( Oryctolagus amiculus ) were shot on farmland in Cambridgeshire, UK over a period of three years. This provided pooled estimates of changes in reproductive and body condition and of fecundity during the annual cycle. Both males and females showed a significant annual cycle in reproductive condition as indicated by changes in the size of the gonads and accessory glands of reproduction. No females were pregnant during October, November and December, but males with active spermatogenesis were found during every month of the year. The distribution of pregnancies through the year showed that the reproductive season varied greatly between individuals. Peak fecundity occurred during April, May and June. On average, each female conceived 23.9 and suckled 17.2 young per breeding season. Proportionately more young died during the early stages of the reproductive season. The body weight of adult males, but not of females, varied seasonally. Body condition (kidney fat index) in adults was maximal at the start of the breeding season and minimal at the end.     

The development of beef breeding bulls

Management of the bull battery will have a dramatic impact on profitability of the cow/calf enterprise. It is critical that young bulls be selected and developed to maximize longevity and productivity for the eventual buyer. Bulls must be structurally sound, healthy, and have adequate libido in order to service the required number of females. Once bulls complete their first breeding season, special care must be taken in order to ensure that they recover and regain needed body condition and pass a bull breeding soundness examination (BBSE). Mature bulls that have reached their genetic potential for growth require less intensive management, but the health program and annual BBSE cannot be overlooked. Mature bulls are also more likely to carry venereal disease and should be screened according to local disease incidence and state regulations. All bulls, regardless of age, should be observed early during the breeding season to ensure that they are physically capable of mounting and servicing females. The establishment of a complete management program, especially for young bulls, is essential to ensure that ranch resources are used efficiently, including maintenance of a high level of reproductive performance of the cow herd.     

Age-related patterns of reproductive success in house sparrows Passer domesticus

A common life history pattern in many organisms is that reproductive success increases with age. We report a similar pattern in house sparrows Passer domesticus , older individuals performed better than yearlings for most measures of reproductive success. Older males and females began breeding earlier in a given season and fledged more young than their yearling counterparts. Individual males also fledged more young in their second breeding season than they did in their first, but individual females did not show consistent improvement in reproductive success from year one to two. A path analysis indicated that age in both sexes acted primarily through the timing of breeding; earlier nesters laid more eggs and hence fledged more young but did not have more nesting attempts. We tested whether the increased reproductive success with age arose from high quality individuals surviving to be older (selection hypothesis). In contrast to the main prediction of this hypothesis that reproductive success and survival should be positively related, we found that survival from one year of age to two years of age was negatively related to reproductive success in the first year for males and females combined. Additionally, individuals that survived to breed as two-year-olds did not differ in total young fledged in their first year from those that did not survive to their second season of breeding. Our results indicate that fledgling production increases with age due to improvements in timing of breeding, particularly in females, and not because of the loss of poor breeders or increased output. Mechanisms producing age-related differences in timing of breeding warrant further study.     

The response of the Great Reed Warbler Acrocephalus arundinaceus to climate change

We examined long-term responses in the breeding performance of the Great Reed Warblers Acrocephalus arundinaceus to climate change. The study took place in various years from 1970 to 2007. During the study period, mean temperatures in the breeding season of the species increased and precipitation decreased significantly. We found evidence for the significant advancement in both earliest and annual median first-egg-laying dates. This advancement correlated with temperature increases early in the season. The latest first-egg laying dates, however, remained unchanged. Other breeding statistics: clutch size, nest losses, and production of young per nest, did not change significantly over the study period. Precipitation did not affect any of the analysed measures. It is important to note, though, that during dry seasons, the production of young per successful nest was higher. In contrast to some woodland species, the Great Reed Warbler seems to adapt well to climate change by shifting laying dates. The reason for this is probably to optimise food resources.     

Conditions at the breeding grounds and migration strategy shape different moult patterns of two populations of Eurasian golden plover Pluvialis apricaria

Events in the life cycle of migrant birds are generally time‐constrained. Moult, together with breeding and migration, is the most energetically demanding annual cycle stages, but it is the only stage that can be scheduled at different times of the year. However, it is still not fully understood what factors determine this scheduling. We compare the timing of primary feather moult in relation to breeding and migration between two populations of Eurasian golden plover Pluvialis apricaria, the continental population breeding in Scandinavia and in N Russia that migrates to the Netherlands and southern Europe, and the Icelandic population that migrates mainly to Ireland and western UK. Moult was studied at the breeding grounds (N Sweden, N Russia, Iceland) and at stopover and wintering sites (S Sweden, the Netherlands). In both populations, primary moult overlapped with incubation and chick rearing, and females started on average 9 d later than males. Icelandic plovers overlapped moult with incubation to a larger extent and stayed in the breeding grounds until primary moult was completed. In contrast, continental birds only moulted the first 5–7 primaries at the breeding grounds and completed moult in stopover and wintering areas, such as S Sweden and the Netherlands. This overlap, although rare in birds, can be understood from an annual cycle perspective. Icelandic plovers presumably need to initiate moult early in the season to be able to complete it at the breeding grounds. The latter is not possible for continental plovers as their breeding season is much shorter due to a harsher climate. Additionally, for this population, moulting all the primaries at the stopover/wintering site is also not possible as too little time would remain to prepare for cold‐spell movements. We conclude that environmental conditions and migration strategy affect the annual scheduling of primary feather moult in the Eurasian golden plover.     

Pouch young removal and return to oestrus in wild southern hairy-nosed wombats (Lasiorhinus latifrons)

The southern hairy-nosed wombat (Lasiorhinus latifrons) is a seasonal breeding, burrowing marsupial adapted to a semi-arid environment and the closest relative of the endangered northern hairy-nosed wombat (Lasiorhinus krefftii). Females typically give birth to one to two young every 3 years with young weaned at 360-400 days. This study examined the occurrence of polyoestry in a wild population of southern hairy-nosed wombats, and in particular the ability of this species to produce additional offspring in the same breeding season if a young was prematurely lost or removed. Pouch young were removed during the breeding seasons of 1996/1997 and 2003. No females from the 1996 (n=3)/1997 (n=3) group gave birth to a second pouch young in the same breeding season. However, two females in this group gave birth to young the following season. In contrast, all the 2003 group of females (n=6) produced a second offspring in the same breeding season after removal of pouch young (RPY). The reason for the different response to RPY between the two groups is unknown. These studies confirm that southern hairy-nosed wombats are polyoestrus in the wild and are capable of producing more than one offspring in a single breeding season. Females that failed to return to oestrus in the breeding season that pouch young were removed bred again in the following season. Rapid replacement of southern hairy-nosed wombat pouch young in the same breeding season as RPY suggests that this procedure, linked to either hand-rearing or interspecific cross-fostering, should be seriously considered as a priority conservation action to increase the population size of the critically endangered sister species, the northern hairy-nosed wombat.     

POPULATION,DIET AND THE ANNUAL CYCLE OF THE LAUGHING DOVE AT BARBERSPAN,PART 4. BREEDING DATA AND POPULATION ESTLMATES

Dean, W. R. J. 1980. Population, diet and the annual cycle of the Laughing Dove at Barber-span, Part 4: Breeding data and population estimates. Ostrich 51:80-91.

From 1974–1976, Laughing Doves Streptopelia senegalensis raised an observed total of 436 young in 619 nest attempts, giving a breeding success of 0,70 young per pair nest attempt, or an overall success of 0,35 young per egg. Breeding success, estimated by computing the probability of survival of an egg through the incubation period, and of a chick through the nestling period suggest that the breeding success is about 0,33 young per egg. Nests were found in every month of the year, though there was a tendency for breeding to be concentrated during the late rainy season and the dry season. The mean annual population size of the Laughing Dove in the study area was estimated at 221 adults and juveniles by one method and at 237 adults and juveniles by another method. The number of young produced each year is correlated with the estimated population size for each year.     

Age-dependent reproductive performance in Northern Goshawks Accipiter gentilis

The age-specific reproductive performance of Northern Goshawks Accipiter gentilis was studied over 22 years in Denmark. The age of the breeding female in relation to the number of young raised was known in 929 breeding attempts, while the age of both the male and the female was known in 496 breeding attempts. The number of fledglings raised per breeding attempt increased with both male and female age, but only for females was it possible to conduct a detailed analysis of this age-dependent relationship. The annual production of fledglings increased with female age from 1 to 7 years of age, whereupon it started to decline. A longitudinal analysis showed that this mean population trend could be attributed to similar age-related trends in individual females. Previous breeding experience did not influence the number of fledglings produced by individual females, and poorly performing females apparently survived with the same probability as well performing ones. The most likely explanation for the age-dependent reproductive performance in the observed Goshawk population appeared to be age-related improvements in competence, such as foraging efficiency.     

The influence of age on reproductive performance in the Brown Thornbill

I examined age effects on reproduction in the Brown Thornbill Acanthiza pusilla in Canberra, Australia. I found that the reproductive performance of both males and females improved with age, although only age-related improvement in male performance had a significant effect on annual reproductive success. Reproductive success improved with male age as a result of improved performance during two stages of the breeding cycle: first-year males were less likely to fledge young than those aged two or more, while both first and second-year males were less successful at raising fledglings to independence than males of three or more. Male performance appears to improve over three years as they gain experience at provisioning nestlings and caring for fledglings without attracting predators, rather than as a direct result of improved foraging skills. In contrast, reproductive success only improved slightly with female age, although females of two or more years initiated their first clutch earlier in the season than one-year-old females, and tended to be more likely to re-nest if a breeding attempt failed. The poor performance of young females appears unlikely to be related to their foraging ability but may be associated with costs imposed by dispersing to a breeding vacancy earlier in the year. Although the reproductive performance of Brown Thornbills improves considerably with age I found no evidence that performance improved as a result of repeated breeding attempts with the same partner.     

POPULATION DYNAMICS OF THE FLIGHTLESS CORMORANT NANNOPTERUM HARRISI

A small population of Flightless Cormorants was followed from 1970 to 1975 inclusive. The birds were extremely sedentary, most never moving more than 2 km from where hatched. Many birds bred several times within a year, almost always with different mates. After successful breeding the mean interval to the next attempt among females was significantly shorter than among males, probably because the male continued to attend the juvenile for longer than did the female. There was an annual peak of nesting in April to November, when sea temperatures were lowest; some nesting occurred in other months but these nests were less successful. About 73% of juveniles survived at least three months after going to sea. Adult females had a significantly higher rate of annual survival (91%) than did males (82%). The mean annual survival of both sexes combined over a 13 year period was 87%. The mean age of first breeding was about 30 months for both males and females. In 1972 breeding success (0·14 young fledged per pair) was much lower than in other years (0·60 young per pair), a lower proportion of juveniles survived, no birds bred for the first time and probably many fewer pairs nested. Adult survival was not affected. This reduced breeding output was associated with an influx of anomalously warm sea water to the area (El Niño). The availability of food is probably both the ultimate and the proximate factor controlling the timing of breeding.     

Spawning periodicity and annual breeding seasons of Tilapia nilotica (L.) in the White Nile

Two annual breeding seasons are indicated for populations of T. nilotica in the White Nile: a major autumnal season (mid-July–September) coinciding with the annual rainfall and a smaller back-up season in February–March. During these seasons reproducing females constituted 80% and 40% of the total population of mature females compared to 10–12% in the intervening summer period. Enhanced gonadal development, fecundity and GSI values characterized the breeding seasons. Gonadal maturation proceeded uninterruptedly even during the summer and repeated fractional spawning is suggested for this species within and beyond the breeding seasons. Most of the spring breeders were either very young or relatively old fish and most medium-sized females bred in the main autumnal season. Older fish were less fecund (by 70–75%) compared to younger fish. In the latter, fecundity increased progressively with body growth (r² = 0.732; p<0.001); the proportionality was lost in medium-sized fish and in older females (0.9–1.5 kg) fecundity correlated negatively with increased body size.     

Density-dependent pup mortality in the Antarctic fur seal Arctocephalus gazellu at South Georgia

The extent and causes of pup mortality in the Antarctic fur seal, Arctocephalus gazella , were investigated at sites of high and low density at South Georgia. Mortality was greater at the high density site, (17—31% of annual pup production) than at the low density site (36%). The main causes of death, starvation and skull injury, occur more frequently at the high density site. Most starvation was caused by failure of the mother-pup bond to form, often caused by disturbance induced by the activities of breeding bulls. In some seasons starvation of older pups may be influenced by reduced food availability but this was usually of minor importance. Injury to the skull resulted from bites inflicted by females either accidently during birth or when pups tried to suck from females other than their mother. Trampling of young pups by bulls was probably responsible for the appreciable incidence of ruptured livers. Infectious disease and drowning played minor roles in pup mortality. Pups born late in the season suffered disproportionately greater mortality which may relate to female age and condition. Food availability (both during and prior to the breeding season) and weather are likely to account for year to year variation in pup mortality rates but the basic rate is primarily determined by breeding density. Further population increase and colonization of new beaches is expected until food resources during the summer, or more probably the winter, become limiting.     

BREEDING PERFORMANCE OF PUFFINS FRATERCULA ARCTICA IN RELATION TO NEST DENSITY, LAYING DATE AND YEAR

The paper presents data on the breeding and predation of Puffins in two areas of different nest density within a single colony on Dun, St Kilda group, Outer Hebrides in 1973-78.
Within a season birds laying early had a slightly higher nesting success than birds laying late, but laying date had little influence on the peak and fledging weights of young. The main disadvantage in late laying was a reduced chance of relaying if the first egg was lost.
Breeding success and chick weights varied from year to year. The 1974 season was the least successful with the lowest nesting success, lowest frequency of feeds, lowest calorific value of feeds, lightest chicks and slowest growth. Overall breeding performance was not related to the annual mean laying dates.
In all years pairs nesting in the area of high nest density did better than pairs nesting at low density. The effect is attributed to differential predation and disturbance by predatory gulls. At least 4.2% of adult Puffins breeding in the area of low burrow density were killed by gulls each breeding season; this is higher than the total annual mortality found in three other studies. Only 0.9% of adults from the high density area were found killed. The subpopulation in the low density area cannot survive without much immigration, yet there is no evidence that this happens.     

Season and testosterone affect contractile properties of fast calling muscles in the gray tree frog Hyla chrysoscelis

In anurans, circulating levels of androgens influence certain secondary sexual characteristics that are expressed only during the breeding season. We studied the contractile properties of external oblique muscles (used to power sound production) in a species of North American gray tree frog, Hyla chrysoscelis, during the breeding season and also in testosterone-treated captive males and females after the breeding season. Compared with the muscles of breeding-season males, the trunk muscles of postbreeding-season males have 50% less mass, 60% longer twitches, and 40% slower shortening velocities. Testosterone levels similar to those found in breeding-season male hylid frogs restore the contractile speed and mass of male trunk muscles and also convert the small slow trunk muscles of females into larger fast-contracting muscles. We conclude that androgens likely play a key role in altering the contractile properties of these muscles in males during the annual cycle, allowing them to operate in the breeding season at the frequencies required to produce the characteristic rapidly pulsed calls of this species. Females as well as nonbreeding-season males do not produce advertising calls, and therefore the slower muscles found in these animals may allow more economic operation of these muscles. The effects of testosterone on female trunk muscles indicate the potential of this hormone in contributing to the sexual dimorphism in size and contractile properties of these muscles, but this dimorphism is likely due to the interaction of more than one hormone.     

Delayed female reproduction in equilibrium and chaotic populations

Behavioural and life history polymorphisms are often observed in animal populations. We analyse the timing of maturation and reproduction in risky and resource-limited environments. Field and laboratory evidence suggests that female voles and mice, for example, can adjust their breeding according to the level of risk to their own survival and to survival probabilities and recruitment of young produced under different environmental conditions. Under risky or harsh conditions breeding can be postponed until later in the current breeding season or even to the next breeding season. We develop a population dynamics and life history model for polymorphism in reproduction (co-existence of breeding and non-breeding behaviours) of females in an age-structured population, with two temporally distinct mating events within the breeding season. We assume that, after overwintering, the females can breed in spring and again in summer or they can delay breeding in spring and breed in summer only. Young females born in spring can either mature and breed in summer or stay immature and postpone breeding over the winter to the next breeding season. We show that an evolutionarily stable breeding strategy is either an age-structured combination of pure breeding behaviours (old females breed and young delay maturity) or a mixed breeding behaviour within age-classes (a fraction of females breed and the rest of the age class postpones breeding). Co-occurrence of mixed reproductive behaviour in spring and summer within a single breeding season is observed in fluctuating populations only. The reproductive patterns depend on intraspecific, possibly interspecific, and ecological factors. The density dependence (e.g. social suppression) and predation risk are shown to be possible evolutionary mechanisms in adjusting the relative proportions of the different but co-existing reproductive behaviours.     

Study of LH response to GnRH in the young male as a criterion of genetic merit for female reproduction in sheep

A high and a low response line in sheep were selected on the basis of the mean concentration of LH in 10-week-old Finn-Dorset ram lambs after an i.v. injection of 5 micrograms GnRH. After 8 male generations the mean LH response of the high line was more than 5-fold that of the low line and the heritability of the selected trait was estimated at 0.44 +/- 0.015. Highly significant line differences in mean LH response to GnRH were also found in males at 20 weeks of age and females at 10 and 20 weeks of age and the genetic correlations between the four LH response traits appear to be close to unity. Large line differences in the mean FSH response to GnRH were also found in both males and females at 10 and 20 weeks of age. Selection had little effect on the physical characteristics of lambs. High-response line ewes entering their first breeding season at about 7 months of age showed oestrus earlier in the season and had higher ovulation rates and numbers of lambs born per ewe lambing than did low-response line ewes. In the second breeding season, at about 19 months of age, the only line difference was a higher ovulation rate early in the breeding season in high-line ewes. It is suggested that these changes may be mediated by a more rapid response in high-line ewes to increased GnRH stimulation at puberty or at the beginning of the breeding season.     

Mate choice in Darwin's Finches

Female Geospiza conirostris on Isla Genovesa, Galapagos, pair preferentially with males who have had previous breeding experience. They choose mates on the basis of courtship behaviour and black adult plumage. By mating with experienced black males, they gain a fitness advantage in terms of fledgling production and recruitment of young into the breeding population. Behavioural signs of past breeding experience and black plumage are reliable age- and condition-dependent traits. We suggest that females use conspicuous black plumage to identify old males at a distance, then interactions through courtship to modify initial assessments. Females paired with inferior males may increase the genetic quality of their offspring by extra-pair copulations; results of heritability analysis of morphology are consistent with this suggestion. Females change mates at a frequency of 12–27% per breeding season. They re-pair with males who are generally old, experienced, and hold territories adjacent to the deserted male. Females that re-pair gain a benefit, whereas males who are deserted within a breeding season incur a cost of more than 50% of their future potential production for that season. We conclude that females in choosing males seek reliable indicators of potential parental care, and in addition they may seek indicators of genetic quality.     

A key factor analysis of a sparrowhawk population

Summary During 1972–86, the Sparrowhawk breeding population in Eskdale, southern Scotland, remained fairly stable, with nest numbers fluctuating between 29 and 39 in different years. The annual production of young varied rather more, between 44 and 86 in different years. In a key factor analysis, the overwinter loss operating in the period between the fledging of young and subsequent recruitment to the breeding population, emerged as the key factor, explaining 77% of the variance in total annual loss, and largely accounting for the pattern of change in breeding numbers. Overwinter loss in each new cohort was also the only form of loss which was density dependent. In a simulation model this loss was found to be capable, on its own, of stabilising the nesting population at the mean level observed. Losses during the breeding season were inversely correlated with losses over the ensuing winter. Thus, good breeding seasons, when many young were produced, were followed by high losses over winter, while poor breeding seasons were followed by reduced losses over winter. No density dependence was detected at any stage of breeding, or in the annual mortality of established breeders.