Fitness consequences of pre- and post-fledging timing decisions in a double-brooded passerine

The fitness consequences of a delayed timing of breeding are expected to affect the temporal characteristics of the whole annual breeding system. One major problem in quantifying the fitness relevance of timing is that individual differences between pairs may cause the seasonal trend. Differentials in juvenile survival due to pre-fledging timing decisions often only appear after fledging of the chicks. Therefore, timing decisions in the post-fledging period, i.e., the duration of parental care, might additionally influence juvenile survival. We tested the effects of timing and parental competence on the post-fledging survival of second-brood juvenile Barn Swallows (Hirundo rustica L.) by swapping earlier and later hatching clutches and radio-tracking the juvenile subjects. The mark-recapture models controlled for the effects of duration of post-fledging care and food availability. There was an annually varying negative seasonal trend in offspring survival that was associated with environmental conditions. Directional selection for early breeding occurred in the two years with scarce autumnal food supply. Furthermore, we found strong selection for long post-fledging parental care. The duration of care neither declined seasonally, nor did longer care compensate for the seasonal decline of juvenile survival. Hence, the reproductive output three weeks after fledging was determined by two parental timing decisions: the timing of breeding and the timing of family breakup. We suggest that differential survival of second-brood fledglings in relation to these decisions is an important part of the selective mechanisms shaping the reproductive system of Barn Swallows.     

Delayed juvenile dispersal and monogamy, but no cooperative breeding in white-breasted mesites (Mesitornis variegata)

Although cooperative breeding is known from only about 9 % of bird species, it has received substantial attention because individuals foregoing their own reproduction to help others represent a long-standing evolutionary puzzle. We studied group formation, breeding system, spatial distribution and several life-history traits of white-breasted mesites (Mesitornis variegata). Based on field observations across 3 years, we found that white-breasted mesites live in year-round stable pairs, and that groups are formed by juvenile philopatry. As other family-living birds, M. variegata exhibit a slow pace-of-life, characterized by high annual adult survival, low productivity, long chick dependence and extended parental care. However, although reproduction is monogamous and juveniles showed interest in their parents’ nests, we found no evidence of cooperative breeding. We suggest that slow life-histories, extended parental care and year-round territoriality predispose juvenile mesites to delay dispersal. However, adult intolerance toward older juveniles may prevent them from adopting a cooperative lifestyle. Comparisons with other species of mesite indicate that monogamy and delayed juvenile dispersal are necessary, but not sufficient for the evolution of cooperative breeding in this family of birds, and that particular ecological and social conditions have facilitated the transition from pair-living to a type of group that may represent a stepping stone in the evolution of cooperative breeding in mesites and other birds.     

Juvenile Survival in a Population of Neotropical Migrant Birds

Determination of population productivity of Neotropical migrant birds and assessment of breeding habitat quality have been based on population densities and nesting success. Data on juvenile survival improve our estimates of population productivity, provide information on factors during the post-fledging period that affect this productivity and, with comparative data, enable us to better assess breeding habitat quality. We present the first estimate of post-fledging juvenile survival in a population of Neotropical migrant birds. We studied post-fledging survival in a population of Wood Thrush ( Hylocichla mustelina) in southern Missouri, (U.S.) an area hypothesized to contain source populations. Nesting success during our study period was 0.266, and individual survival within the nest was 0.245. Post-fledging survival during the first 8 weeks after fledging was 0.423. Survival varied significantly between post-fledging weekly age classes, with survival of weeks 1, 2, 3, and 4 through 8 being 0.716, 0.930, 0.637, and 1.00, respectively. Probability of predation after fledging was 0.506. Probability of mortality by other causes was 0.071. Probability of predation varied by weekly age class and may have been related to behaviors occurring at different developmental stages. Post-fledging survival was not correlated with nestling mass and did not change throughout the course of the breeding season. Analysis of the source/sink status of the population based on our estimates of nesting success and post-fledging survival indicates that young were being produced below replacement levels during our study period. Large-scale management decisions should take into account potential fluctuations in the productivity of Neotropical migrant populations over time. Data on post-fledging juvenile survival are needed from other populations of Neotropical migrant birds to more accurately assess differential productivity between populations and better assess breeding habitat quality.     

Estimating annual survival in sexually dimorphic species from proportions of first-year birds

We estimated the annual adult survival (S) of several species of temperate and tropical sexually dimorphic tanagers and manakins from the proportions of first-year and older individuals in museum collections. In the case of the sexually dimorphic species included here, young males through their first year resemble females and can be distinguished from older males. We assumed that the sex ratio among first-year birds is even and estimated the number of immature females (IF) in a sample as equal to the number of immature males (IM), using sex information on museum labels. The number of adult females (AF) is the total number of females (F) minus IF, and survival can be estimated by S = AF/F. We used a Monte Carlo resampling approach to estimate standard errors and 1% and 99% confidence limits on S for samples of size N, based on 1000 sets of N randomly drawn individuals and observed proportions of the F, IM, and AM classes. Estimates of annual survival (S) were 0.60-0.73 in four North American species of Piranga tanagers and 0.50 and 0.62 in two montane Central American species. Values of S in lowland species of Habia and Ramphocelus tanagers ranged between 0.68 and 0.82. Annual survival in Pipra manakins was 0.53-0.68, and values for two Manacus species were 0.69 and 0.70. Where estimates of apparent survival (phi) from capture-recapture studies were available, the two methods produced correlated results with a tendency for phi < S. Among tanagers, only Ramphocelus dimidiatus (0.82) and Piranga bidentata (0.50) stood out from the other species, for which S averaged 0.66 +/- 0.05 (SD). Annual survival rates of temperate and tropical species overlapped broadly in this analysis, reinforcing the conclusion that distinctive life-history traits of temperate vs. tropical species cannot be explained completely by the adjustment of reproductive investment in relation to adult survival and the expectation of future life.     

Modeling post-fledging survival of Lark Buntings in response to ecological and biological factors

We evaluated the influences of several ecological, biological, and methodological factors on post-fledging survival of a shortgrass prairie bird, the Lark Bunting (Calamospiza melanocorys). We estimated daily post-fledging survival (n = 206, 82 broods) using radiotelemetry and color bands to track fledglings. Daily survival probabilities were best explained by drought intensity, time in season (quadratic trend), ages < or = 3 d post-fledging, and rank given drought intensity. Drought intensity had a strong negative effect on survival. Rank was an important predictor of fledgling survival only during the severe drought of 2002 when the smallest fledglings had lower survival. Recently fledged young (ages < or = 3 d post-fledging) undergoing the transition from nest to surrounding habitat experienced markedly lower survival, demonstrating the vulnerable nature of this time period. Survival was greater in mid and late season than early season, corresponding to our assumptions of food availability. Neither mark type nor sex of attending parent influenced survival. The model-averaged product of the 22-d survival calculated using mean rank and median value of time in season was 0.360 +/- 0.08 in 2001 and 0.276 +/- 0.08 in 2002. Survival estimates that account for age, condition of young, ecological conditions, and other factors are important for parameterization of realistic population models. Biologists using population growth models to elucidate mechanisms of population declines should attempt to estimate species-specific of post-fledging survival rather than use generalized estimates.     

What are the strengths and limitations of direct and indirect assessment of dispersal? Insights from a long-term field study in a group-living bird species

Molecular methods of assessing dispersal have become increasingly powerful and have superseded direct methods of studying dispersal. Although now less popular, direct methods of studying dispersal remain important tools for understanding the evolution of dispersal. Here, we use data from Siberian jays Perisoreus infaustus, a group-living bird species, to compare natal dispersal distances and rates using visual mark–recapture, radio-tracking and microsatellite data. Siberian jays have bimodal natal dispersal timing; socially dominant offspring remain with their parents for up to 5 years (delayed dispersers), while they force their subordinate brood mates to leave the parental territory at independence (early dispersers). Early dispersers moved about 9,000 m (visual mark–recapture, radio-tracking) before settling in a group as a non-breeder. In contrast, delayed dispersers moved about 1,250 m (visual mark–recapture only) and mainly moved to a breeding opening. Dispersal distances were greater in managed habitat compared to natural habitat for both early and delayed dispersers. Molecular estimates based on 23 microsatellite loci and geographical locations supported distance estimates from the direct methods. Our study shows that molecular methods are at least 22 times cheaper than direct methods and match estimates of dispersal distance from direct methods. However, molecular estimates do not give insight into the behavioural mechanisms behind dispersal decisions. Thus, to understand the evolution of dispersal, it is important to combine direct and indirect methods, which will give insights into the behavioural processes affecting dispersal decisions, allowing proximate dispersal decisions to be linked to the ultimate consequences thereof.     

Fitness costs of dispersal in red foxes (Vulpes vulpes)

The costs of dispersal are an important factor promoting natal philopatry, thereby encouraging the formation of social groups. The red fox, Vulpes vulpes, exhibits a highly flexible social system and one that is thought to represent a possible stage in the evolution of more complex patterns of group-living. Although the potential benefits accruing to philopatric offspring have previously been studied in this species, the potential costs of dispersal have received less attention. We contrasted survival rates, nutritional status, injuries and reproductive output of dispersing and non-dispersing male and female foxes in an urban population to assess the relative costs of dispersal versus natal philopatry. Mortality rates were not significantly higher for dispersing foxes, either in the short- or long-term. There was no evidence of increased nutritional stress in dispersing individuals. Dispersing individuals did, however, exhibit greater levels of wounding, although this did not appear to affect survival. Dispersing females were more likely to miss a breeding opportunity early in their reproductive lifespan. In contrast, both dispersing and non-dispersing males were unlikely to breed in their first year. We conclude that the major fitness component in females affected by dispersing is age at first reproduction.     

Survival probability estimates for the endangered loggerhead sea turtle resident in southern Great Barrier Reef waters

Sex- and age-class-specific survival of a loggerhead turtle population resident in southern Great Barrier Reef waters was estimated using a long-term capture-mark-recapture (CMR) study and the Cormack-Jolly-Seber modelling approach. The CMR history profiles for 271 loggerheads tagged over 9 years (1984-1992) were classified into two age classes (adult, immature) based on somatic growth and reproductive traits. The sex and maturity status of each turtle was determined from visual examination of reproductive organs using laparoscopy. A reduced-parameter model accounting for constant survival with sex- and time-specific recapture was adequate for estimating age-class-specific survival probabilities, but inclusion of time-specific transient behaviour was informative for the immature age class. The annual fluctuations in the estimated proportion of transient immatures was not a function of sampling effort, but could be due to anomalous oceanographic conditions affecting dispersal of the immature class. There was no sex-specific difference in survival probabilities for either age class, but females were more likely to be recaptured than males, which might be related to behavioural differences such as sex-biased dispersal. The expected annual survival probability for adults was 0.875 (95% CI: 0.84-0.91). The expected annual survival probability for immatures was 0.859 (95% CI: 0.83-0.89), but when the transients were accounted for, the expected annual survival for the resident immature loggerheads was 0.918 (95% CI: 0.88-0.96). These are the first substantive estimates of annual survival probabilities for any loggerhead sea-turtle stock and provide a basis for developing a better understanding of loggerhead population dynamics.     

Potential effects of environmental contamination on Yuma Myotis demography and population growth.

Unplanned natural and anthropogenic disasters provide unique opportunities for investigating the influence of perturbations on population vital rates and species recovery times. We investigated the potential effects of a major pesticide spill by comparing annual survival rates using mark-recapture techniques on a riparian bat species, Yuma Myotis (Myotis yumanensis). Demography and population dynamics for most bat species remain poorly understood despite advances in mark-recapture estimation and modeling techniques. We compared survival and population growth rates of two roost populations exposed to a large chemical (metam sodium) spill in the upper Sacramento River in Northern California with two roost populations outside the contaminated area from 1992 to 1996. Hypotheses about long-term effects of the spill on female juvenile and adult survival were tested using an information-theoretic approach (AIC). Working hypotheses included effects of age, chemical spill, and time trend on survival. Female adult survival was higher than female juvenile survival across all sites, suggesting stage-specific mortality risks. Model-averaged estimates of female juvenile survival in the contaminated area (0.50-0.74) were lower than in control roosts (0.60-0.78) for each year in the study, suggesting that the spill may have reduced juvenile survival for several years. Female adult survival (0.73-0.89) did not appear to be strongly affected by the spill during the years of the study. There was an increase in survival for both stage-classes across all populations during the study period, which may have been caused by the end of an extended drought in California in the winter of 1993. The spill-affected population was in decline for the first year of the study as indicated by an estimated growth rate (lambda) < 1, but population growth rates increased during the four-year period.     

Transition to independence and evidence of extended parental care in the gentoo penguin (<Emphasis Type="Italic">Pygoscelis papua</Emphasis>)

We used radio telemetry and observations to study the activity patterns and behavior of gentoo penguin chicks at Admiralty Bay, King George Island, South Shetland Islands in 2005 during their “fledging period”; defined as the time between a chick’s first trip to sea and its final dispersal from the breeding colony. Gentoo penguins exhibited delayed dispersal of young and extended parental provisioning, behaviors not observed in other Pygoscelis species. Chicks took their first trip to sea at a mean age of 70 days of age, before finally departing the colony at a mean age of 82 days. During this fledging period, individual chicks made an average of five trips to sea. Trip duration increased significantly as chicks aged, with trips to sea becoming similar to literature values of adult foraging trips in both timing and duration. Behavioral observations and mass dynamics confirmed that many chicks were still being fed during this fledging period, with parental feeding behaviors most often observed in the late afternoon to evening hours. We hypothesize that these behaviors provide an opportunity for chicks to gain experience at sea prior to dispersal and might allow them to develop foraging skills before they are completely independent.     

Sex-specific effects of the local social environment on juvenile post-fledging dispersal in great tits

An individual’s decision to disperse from the natal habitat can affect its future fitness prospects. Especially in species with sex-biased dispersal, we expect the cost–benefit balance for dispersal to vary according to the social environment (e.g., local sex ratio and density). However, little is known about the social factors affecting dispersal decisions and about the temporal and spatial patterns of the dispersal process. In our study, we investigated experimentally the effects of the social environment on post-fledging dispersal of juvenile great tits by simultaneously manipulating the density and sex ratio of fledglings within forest plots. We expected young females in the post-fledging period mainly to compete for resources related to food and, as they are subordinate to males, we predicted higher female dispersal from male-biased plots. Juvenile males compete for vacant territories already in late summer and autumn; thus, we predicted increased male dispersal from high density and male-biased plots. We found that juvenile females had a higher probability to leave male-biased plots and had dispersed further from male-biased plots in the later post-fledging phase when juvenile males start to become territorial and more aggressive. Juvenile males were least likely to leave male-biased plots and had smallest dispersal distances from female-biased plots early after fledging. The results suggest that the social environment differentially affected the costs and benefits of philopatry for male and female juveniles. The local sex ratio of individuals is thus an important social trait to be considered for understanding sex-specific dispersal processes.     

Estimating species-specific survival and movement when species identification is uncertain

Incorporating uncertainty in the investigation of ecological studies has been the topic of an increasing body of research. In particular, mark-recapture methodology has shown that incorporating uncertainty in the probability of detecting individuals in populations enables accurate estimation of population-level processes such as survival, reproduction, and dispersal. Recent advances in mark-recapture methodology have included estimating population-level processes for biologically important groups despite the misassignment of individuals to those groups. Examples include estimating rates of apparent survival despite less than perfect accuracy when identifying individuals to gender or breeding state. Here we introduce a method for estimating apparent survival and dispersal in species that co-occur but that are difficult to distinguish. We use data from co-occurring populations of meadow voles (Microtus pennsylvanicus) and montane voles (M. montanus) in addition to simulated data to show that ignoring species uncertainty can lead to biased estimates of population processes. The incorporation of species uncertainty in mark-recapture studies should aid future research investigating ecological concepts such as interspecific competition, niche differentiation, and spatial population dynamics in sibling species.     

The evolution of marmot sociality: I. Why disperse late?

Summary Prolonged toleration of offspring in marmots was hypothesized to be (1) a means of preventing dispersal of undersized young (Barash 1974 a) or more generally (2) continued parental investment, increasing the probability of descendant survival and reproduction (Armitage 1981, 1987). These hypotheses are tested in this paper for one of the most social of marmot species, the alpine marmot. The animals studied lived in groups within territories defended by a dominant male and female, or as floaters lacking a well-defined home range. Offspring did not disperse before sexual maturity at age 2 (Fig. 1). Only territorial females bred, whereas territorial males were not able to monopolize reproduction likewise (Table 2). Dispersers had similar spring mass to nondispersers (Table 4). Hence, hypothesis 1 is not supported, at least not for adult-sized, > 2 years old animals. During their residency, 19% of subordinates obtained their natal territory or a neighboring one (Fig. 2). Long distance dispersal bore a high mortality risk. Thus, toleration of mature offspring could well represent parental investment. Other results, however, suggest additional influences on the timing of dispersal. (i) Males dispersed later than females (Fig. 3), possibly because of mate sharing by territorial males (see Emlen 1982). (ii) The higher mass loss of dispersers during the previous winter indicates that weak animals were forced to leave (Table 5) despite presumably lower chances of becoming territorial (Table 3). (iii) Subordinate animals which could not be the offspring of both territorials present were not more likely to disperse (Fig. 3). (iv) Lower dispersal rates when immatures lived in the group (Fig. 3) may indicate benefits from the subordinates' presence for rearing young.     

Offspring sex ratio in relation to parental structural size and body condition in the long-lived wandering albatross (<Emphasis Type="Italic">Diomedea exulans</Emphasis>)

Sex ratio theory is one of the most controversial topics in evolutionary ecology. Many deviations from an equal production of males and females are reported in the literature, but few patterns appear to hold across species or populations. There is clearly a need to identify fitness effects of sex ratio variation. We studied this aspect in a population of a long-lived seabird, the wandering albatross (Diomedea exulans), using molecular sex-identification techniques. We report that parental traits affect both (1) fledgling traits in a sex-dependent way and (2) chick sex: Sons are overproduced when likely to be large at fledging and, to a lesser extent, daughters are overproduced when likely to be in good body condition at fledging. Because for the same population, a previous study reported that post-fledging survival was positively affected by size in males and by body condition in females, our results suggest that wandering albatrosses manipulate offspring sex to increase post-fledging survival.     

Roles of survival and dispersal in reintroduction success of Griffon vulture (Gyps fulvus).

The success of reintroduction programs greatly depends on the amount of mortality and dispersal of the released individuals. Although local environmental pressures are likely to play an important role in these processes, they have rarely been investigated because of the lack of spatial replicates of reintroduction. In the present study, we analyzed a 25-year data set encompassing 272 individuals released in five reintroduction programs of Griffon Vultures (Gyps fulvus) in France to examine the respective roles of survival and dispersal in program successes and failures. We use recent developments in multi-strata capture-recapture models to take into account tag loss in survival estimates and to consider and estimate dispersal among release areas. We also examined the effects of sex, age, time, area, and release status on survival, and we tested whether dispersal patterns among release areas were consistent with habitat selection theories. Results indicated that the survival of released adults was reduced during the first year after release, with no difference between sexes. Taking into account local observations only, we found that early survival rates varied across sites. However when we distinguished dispersal from mortality, early survival rates became equal across release sites. It thus appears that among reintroduction programs difference in failure and success was due to differential dispersal among release sites. We revealed asymmetrical patterns of dispersal due to conspecific attraction: dispersers selected the closest and the largest population. We showed that mortality can be homogeneous from one program to another while, on the contrary, dispersal is highly dependent on the matrix of established populations. Dispersal behavior is thus of major interest for metapopulation restoration and should be taken into account in planning reintroduction designs.     

Increasing the Accuracy of Productivity and Survival Estimates in Assessing Landbird Population Status

Abstract:  The conservation of species with declining populations requires information on population demography and identification of factors that limit population growth. For landbird species, an understanding of large-scale population declines often requires assessment of local population processes, including the production of offspring, the survival of those offspring, and adult survival. Population growth has been modeled for several species of landbirds to date, and these studies have provided important information on relationships between population status and population-limiting factors. Several recent studies have illuminated field methods and analytical techniques that can aid in increasing the accuracy of productivity and survival estimates for population models. We reviewed these methods and recommend their implementation, including quantification of the season-long productivity of individuals, collection of empirical data on juvenile survival during the postfledging and overwintering periods, and incorporation of adult breeding dispersal into annual adult survival estimates. Such methods will allow for more accurate assessment of population status and provide a better understanding of the factors on which to focus our conservation efforts.     

Population biology of White-crowned Sparrows: Residence time and local movements of juveniles

Summary A field study of the ranging behavior of post-fledging White-crowned Sparrows (Zonotrichia leucophrys nuttalli) was carried out in undisturbed natural habitat in the Point Reyes National Seashore, Marin Co., California, USA. Forty-three first-brood juveniles, with tail feathers still growing, were banded at an estimated average age of 27 days and re-trapped an average of an additional 24 days for a total minimum time in residence in the immediate natal area of 51 days (Table 1).The distance from the site of first capture to subsequent recapture sites was about 60 m until age 35 days; then the distance increased to about 250 m by age 50 days (Fig. 2). These data indicate that the average juvenile White-crowned Sparrow completes at least the first 50 days of life in its natal area. Experimental work on song ontogeny has previously demonstrated that the sensitive period for song learning ends at about 50 days. Hence, young in our study population probably learn their natal dialect prior to any major dispersal movements. The correspondence between residence time and song learning would have the consequence of perpetuating song dialects as geographically structured populations.     

Great spotted cuckoo fledglings are disadvantaged by magpie host parents when reared together with magpie nestlings

The post-fledging period is a critical phase for juvenile survival, and parental care provided during this period is a key component of avian reproductive performance. Very little is known about the relationships between foster parents and fledglings of brood parasites. Here, we present the results of a 5-year study about the relationships between fledglings of the non-evictor brood parasitic great spotted cuckoo (Clamator glandarius) and its magpie (Pica pica) foster parents. Sometimes, great spotted cuckoo and magpie nestlings from the same nest can fledge successfully, but most often parasitic nestlings outcompete host nestlings and only cuckoos leave the nest. We have studied several aspects of cuckoo post-fledging performance (i.e. feeding behaviour, parental defence and fledgling survival) in experimental nests in which only cuckoos or both magpie and cuckoo nestlings survived until leaving the nest. The results indicate that great spotted cuckoo fledglings reared in mixed broods together with magpie nestlings were disadvantaged by magpie adults with respect to feeding patterns. Fledgling cuckoos reared in mixed broods were fed less frequently than those reared in only cuckoo broods, and magpie adults approached less frequently to feed cuckoos from mixed broods than cuckoos from only cuckoo broods. These results imply that the presence of host's own nestlings for comparison may be a crucial clue favouring the evolution of fledgling discrimination; and furthermore, that the risk of discrimination at the fledgling stage probably is an important selection pressure driving the evolution of the arms race between brood parasites and their hosts.     

Allozymes and morphometric characters of three species ofMytilus in the Northern and Southern Hemispheres

Many authors have considered the common mussels in temperate waters of the Northern and Southern Hemispheres to be a single cosmopolitan species,Mytilus edulis Linnaeus, 1758. Others have divided these mussels into several subspecies or species. Samples of mussels were collected from 36 locations in the Northern Hemisphere and nine locations in the Southern Hemisphere. Electrophoretic evidence from eight loci indicates that the Northern Hemisphere samples consist of three electrophoretically distinguishable species:M. edulis from eastern North America and western Europe;M. galloprovincialis Lamarck, 1819 from the Mediterranean Sea, western Europe, California, and eastern Asia; andM. trossulus Gould, 1850 from the Baltic Sea, eastern Canada, western North America and the Pacific coast of Siberia. Mussels from Chile, Argentina, the Falkland Islands and the Kerguelen Islands contain alleles characteristic of all three Northern Hemisphere species, but because they are most similar toM. edulis from the Northern Hemisphere, we suggest that they tentatively be included inM. edulis. These South American samples are morphologically intermediate between Northern HemisphereM. edulis andM. trossulus. Mussels from Australia and New Zealand are similar in allele frequency and morphometric characters toM. galloprovincialis from the Northern Hemisphere. FossilMytilus sp. are present in Australia, New Zealand and South America, which suggests that the Southern Hemisphere populations may be native, rather than introduced by humans. Morphometric characters were measured on samples which the allozyme data indicated contained a single species. Canonical variates analysis of the morphometric characters yields functions which distinguish among our samples of the species in the Northern Hemisphere.