Effects of Aging and Mate Retention on Reproductive Success of Captive Female Peregrine Falcons

SYNOPSIS. Separating ecological (extrinsic) factors affectingreproduction from physiological and experiential (intrinsic)factors can be problematic in free-ranging animals. This studyexamined age-related changes in six measures of reproductivesuccess (clutch size, fertility, hatchability, brood size athatching, survivorship of nestlings, and brood size at fledging)for captive female peregrine falcons (Falco peregrinus) whereecological factors (i.e., access to mates, nesting sites, andfood) were not limiting and were similar across years and betweenindividuals. Mean nestling survivorship increased throughoutthe lifespan of the female while all other measures of reproductionpeaked at about seven years of age and decreased thereafter.Birds with prior breeding experience had higher productivitythan inexperienced birds of the same age. Productivity increasedwith increasing experience of the pair. Productivity droppedby an average of 53% when females experienced a change of mate,and then subsequently increased over a period of several years.Productivity was not affected when birds were moved to differentbreeding sites. There was no evidence that early improvementsin reproduction were related to chronological aging in the absenceof experiential differences. Birds that began breeding earlierproduced more fledglings during their lifetimes as a resultof higher annual productivity. Lifetime reproduction was notcorrelated with longevity because birds with higher maximalegg production had shorter breeding lifespans. Birds retainingmates produced more fledglings during their lifetime than birdsthat changed mates, but birds that changed mates more than oncedid not have lower lifetime reproduction than birds that hadonly one mate change. These results suggest that 1) age-relatedchanges in reproduction are not necessarily resource-mediated,2) in the absence of resource limitation, experience of thepair is a primary factor determining annual reproductive success,3) benefits of increasing experience may be offset by the onsetof senescence, 4) the cost of present egg production on futurereproductive potential supports a "pleiotropic" theory of aging,and 5) costs associated with mate changes may encourage selectionfor low "divorce" rates (i.e., lifetime monogamy) in this species.     

Intermittent breeding in Cory's Shearwater Calonectris diomedea of Selvagem Grande, North Atlantic

Every year, 10% of the breeding population of Cory's Shearwater Calonectris diomedea of Selvagem Grande, between the Canaries and Madeira (30̀09'N, 15̀52'W), fail to breed. These "sabbatical" periods can last up to 7 years but typically last 1 year in 58% of the cases, with an average of 1.7 years. Both sexes are involved, and most are young breeding birds. The frequency and duration of the absences decrease with the increase of breeding experience. During the year preceding the absence, the breeding success, mate fidelity and nest site fidelity of the breeding birds are the same as those of their sabbatical congeners. During the sabbatical absence, the former mate is also absent in just less than half of the cases. Of those present, two-thirds are at the same nest site and one-third at another nest site. On the return of the "missing" birds, the fidelity both to the former mate and the former nest site is low, and more so if the length of the absence has been over 1 year, because the former mate and the former nest site are then seldom available. Divorce is frequent and the breeding success is low. Comparisons with other Procellariiformes reveal few differences in the intermittent breeding except for the proportions involved, which are six times greater for some species than for others.     

Reproductive consequences of natal dispersal in a highly philopatric seabird

Natal and breeding dispersal have a major impact on gene flowand population structure. We examined the consequences of nataldispersal on the reproductive success (proportion of pairs rearingchicks) of colonial-breeding Thick-billed murres (Uria lomvia).Reproductive success increased with distance dispersed for thefirst and second breeding attempt. The increase in breedingsuccess leveled off at natal dispersal distances above 7 m.Our results were consistent with the idea that the relationshipbetween dispersal and reproductive success is caused by siteavailability and mate choice as birds willing to disperse fartherhad a greater choice of potential sites and mates. This hypothesiswas supported by the fact that birds dispersing farther weremore likely to pair with an experienced breeder, which increasesthe likelihood of breeding success for young breeders. Explanationsfor increasing breeding success with increased dispersal basedon inbreeding effects were unlikely because most breeding failureswere caused by egg loss rather than infertility or nestlingdeath. However, we could not explain why >50% of birds returnwithin 3 m of the natal site, despite having an up to 50% lowerreproductive success than birds dispersing 7 m or more.     

High divorce rates in Corsican blue tits: how to choose a better option in a harsh environment

We investigate which hypothesis, the “better mate hypothesis” or the “better territory hypothesis” best explains the unusually high divorce rate (59%) in a population of blue tits (Parus caeruleus) living in a sclerophyllous habitat characterised by severe environmental constraints (trophic, parasitic, climatic) on the island of Corsica, France. Using data from the breeding seasons 1985–1998 and from a brood size experiment (1990–1993) we examined the causes of divorce and their consequences on breeding performance, mate assortment and territory choice. Breeding performance had no significant effect on whether birds re‐united or divorced in the next breeding season. Re‐uniting pairs did better than divorced females and the latter improved their breeding performance compared to prior to divorce, but this was mainly due to age and territory effects. There were no differences in male performance depending on whether they re‐united or divorced. The age combination of pairs did not differ between re‐uniting and divorcing pairs, but mate assortment changed after divorce with males re‐mating more often with older partners than females. Manipulation of brood size showed a trend for birds with enlarged broods to divorce more. Pairs responded significantly to territory quality by divorcing more often in poor than in good breeding sites. Both faithful pairs and male divorcees had shorter breeding dispersal distances than female divorcees. Divorce rates were determined by the large differences in quality among breeding sites. Males, whatever their status, usually retained their previous territory whereas divorced females moved significantly longer distances and improved their breeding site. Moving to a better territory after divorce benefits only females which appear to be the choosing sex in the decision to divorce. This study strongly supports the “habitat mediated hypothesis” and we suggest that the large observed intraspecific variation in the magnitude of divorce rates in many species of birds is mostly determined by habitat characteristics.     

Pair bond and breeding success in Blue Tits Parus caeruleus and Great Tits Parus major

Data from 939 nests of the Blue Tit Parus caeruleus and 1008 nests of the Great Tit P. major from nestboxes provided in superabundance in mixed forest study sites between 1976 and 2001 were analysed to examine the effects of mate retention on breeding success and the relationship between mate fidelity and site fidelity. Most birds retained their former partner (76% in Great Tits and 65% in Blue Tits). The probability of a pair divorcing was affected by male age in Great Tits, divorce being more likely in pairs with first‐year males. Great Tit pairs breeding together for a second season bred earlier, but had no higher breeding success than pairs breeding together for the first time. In Blue Tits laying date and start of incubation tended to be earlier in pairs breeding together for a second season, but hatching and fledging dates were not earlier than in other pairs. Great Tit pairs breeding together for two consecutive seasons bred earlier in the second season than in the first, but breeding success did not differ significantly between years. In both species, breeding performance did not differ between pairs that divorced after a season and pairs that stayed together. Thus breeding success did not determine whether a pair divorced or bred together again. Neither Blue Tits nor Great Tits improved their breeding performance through divorce. Blue Tit females even had fewer fledglings in the year after divorce than in the year before. Mate retention affected breeding site fidelity. Blue Tit females had greater breeding dispersal distances between consecutive years when re‐mating than when breeding again with the same mate. In Great Tits both males and females dispersed more when re‐mating than when retaining the former partner, suggesting that mate retention increased the chance of retaining the breeding site. In both species, breeding dispersal distances did not differ between pairs that divorced and pairs in which one mate disappeared. Because no major advantage of mate retention was evident, we suggest that mate retention evolved under different conditions than those found in study sites with high breeding densities and a superabundance of artificial nesting sites.     

Within-season divorce in Blue Tits (Cyanistes caeruleus)

Divorce between breeding seasons, i.e. mate change while the old breeding partner is still alive, has been well-studied in several bird species. It can be viewed either as the result of reproductive decisions of individuals to maximize their own fitness or as the outcome of intra-sexual competition for mating opportunities. In contrast, divorce within a breeding season—also referred to as rapid mate switching—has received much less attention. Within-season divorce may occur after sudden changes in environmental conditions, such as those causing the disappearance or devaluation of nesting sites or territories, or when better breeding partners become available. Within-season divorce can be initiated by a member of the pair, or it can be the result of a take-over by an unpaired individual that competes for access to the resource. During a field experiment investigating the effects of limiting nesting sites on reproductive behaviour in Blue Tits, we recorded several cases of within-season divorce. The rate of divorce was not related to the experimental nest-site limitation, and pairs that changed their partner suffered reduced reproductive success compared to faithful pairs. Although there were no differences in the timing of breeding, clutch size or hatching success, pairs with a new partner also suffered a reduced fledging success, which was partly explained by complete brood failures. This study highlights that the pair bond prior to egg laying can be unstable when conditions force individuals to compete for a new partner or nest site and indicates the importance of the correct timing of divorce within the breeding cycle.     

Is clutch size individually optimized?

Brood size manipulations were carried out to test whether clutchsize variation in individual great tits (Parus major) controlledfor laying date was tuned to their phenotypic quality and/orlocal food abundance (individual optimization hypothesis; IOH).Broods with different original clutch sizes, but equal hatchingdates, were manipulated to a common brood size. A third broodwas kept as a control. Under the IOH, we expected a positiveassociation between reproductive success and original clutchsize. Fledgling production varied in an inconclusive way aftermanipulation, with data from 1 out of 3 years favoring the IOH.The effect of manipulation on the probability of a second clutchwas consistent with the IOH in another 1 out of the 3 years.When fitness accrued to second broods was also taken into considerationin terms of annual fledgling production, results from 2 outof 3 years tended to support the IOH. There was no effect ofthe manipulation on fitness (estimated as the number of recruitsplus parents breeding in the next season). Both the clutch component(local recruitment) and the parental component (survival tillnext breeding season) varied inconclusively with respect tothe IOH. On the basis of fitness measurements, the IOH couldnot be confirmed as an explanation for clutch size variationin this population. In 2 out of 3 years one of the three fitnesscomponents measured varied in accordance with the IOH. Overallthe evidence for the IOH in this data set is therefore weak.     

Mate fidelity and breeding site tenacity in a monogamous sandpiper, the black turnstone

We examined the relationship between mate fidelity and breeding site tenacity during a 5-year study of the black turnstone, Arenaria melanocephala, a socially monogamous sandpiper breeding in subArctic Alaska. We tested the predictions of several hypotheses regarding the incidence of divorce and the benefits of fidelity to mate and breeding site. Interannual return rates to the breeding grounds (88% for males, 79% for females) were among the highest yet recorded for any scolopacid sandpiper, and 88% of returning birds nested on their previous year's territory. The annual divorce rate was only 11%, and mate fidelity was significantly linked to fidelity to territory but independent of sex and year. Males arrived in spring significantly earlier than their mates and interannual fidelity was influenced by the relative timing of arrival of pair members. Reunited pairs had significantly higher fledging success than new pairs formed after death or divorce. The incidence of divorce was unrelated to reproductive success the previous year, although birds nested significantly further away after failure than after a successful nesting attempt. Sightings of marked individuals suggested that members of pairs do not winter together, and breeding site tenacity provides a mechanism through which pair members can reunite. We reject the 'incompatibility' hypothesis for divorce in turnstones, and our data contradict predictions of the 'better option' hypothesis. Alternatively, we propose the 'bet-hedging' hypothesis to explain the occurrence of divorce, which transpires when an individual pairs with a new mate to avoid the cost of waiting for a previous mate to return. Such costs can include remaining unmated, if the former mate has died, or experiencing lower reproductive success because of delayed breeding. Copyright 2000 The Association for the Study of Animal Behaviour.     

Breeding performance,mate fidelity,and nest site fidelity in a long‐lived seabird: behaving against the current?

There is evidence that breeding failure is associated with divorce and dispersal in many bird species. However, deviations from the general pattern “success‐stay/failure‐leave” seem to be common, suggesting that factors other than breeding performance may importantly influence mate and habitat selection. Moreover, variability in response to performance suggests coexistence of different evolutionary strategies of mate and site selection within a population. In this study, we assessed how individuals conform to the success‐stay/failure‐leave pattern in kittiwakes (Rissa tridactyla), and aimed to identify categories of individuals presenting different behavioural patterns. We considered individual attributes (experience, prior residence at the nest site, performance in multiple breeding attempts), pair attributes (arrival asynchrony, timing of failure, pair duration), and productivity in habitat patches. Timing of failure was an important factor. Pair reunion probability was close to 0.5 in failed pairs, but it was consistently higher in early failed than in late failed pairs. Prior residence better explained variability in probability of reunion in failed pairs than pair duration. However, the positive influence of prior residence on the probability of reunion was perceptible only in early failed pairs. Divorce probability in successful pairs increased with arrival asynchrony, and was higher in first‐time than in experienced breeders. Local productivity positively influenced site fidelity probability in early failed birds, but not in late failed ones. Using memory models, we found that dispersal decisions integrate information on individual breeding performance in a temporal scale longer than one year. This study contributed to the identification of relevant states to be considered when addressing mate and nest site choice. Natural selection may operate on slight fitness differences that cannot be detected without high levels of stratification according to the appropriate individual and habitat attributes.     

The effects of sex, age and breeding success on breeding dispersal of pied flycatchers along a pollution gradient

We modelled breeding dispersal of an insectivorous bird, the pied flycatcher (Ficedula hypoleuca) around a point source of heavy metals (a copper smelter). We tested for the effects of sex, age, breeding success and environmental pollution on breeding dispersal distances of F. hypoleuca females and males. Unlike many earlier studies on breeding dispersal, we took into account distance-dependent sampling bias by including in our model the recapture probabilities at different distances from the site of origin. Our results show that F. hypoleuca females disperse much farther (on average 7.9 km) from their breeding sites than what was previously thought. In contrast, males only disperse short distances (on average 190 m). Breeding success affected female breeding dispersal distances depending on female age: young females moved on average 8 km from their previous breeding place irrespective of their breeding success, while old females only seemed to move this far when their fledgling production was good. Females successful in their breeding dispersed as far as less successful females, or, among old birds, even farther. Females which dispersed long distances produced more fledglings after the movement than those staying near their previous breeding site. Degree of environmental pollution had no effect on female or male breeding dispersal distances. A polluted and unproductive environment does not seem to stimulate F. hypoleuca parents to move to more profitable territories.     

Effects of site fidelity and breeding performance on mate retention in a short‐lived passerine,the tree swallow Thachycineta bicolor

We studied the relationship among re‐mating, site fidelity and breeding performance in the tree swallow Tachycineta bicolor using 16 y of data on reproductive biology in a population breeding in nest boxes near Ithaca, New York. Of 217 pairs for which both members survived the non‐breeding season, 76% mated with a new partner and 24% reunited with their previous mate. Pairs did not increase their breeding success by breeding together for more than one breeding season. Males produced fewer fledglings after breeding with a new partner, but females neither increased nor decreased their success when breeding with a new mate. Females who bred with a new partner were younger than females that reunited with their previous mates, and they were more likely to move to a different nest box. Males that bred with a new mate were of similar age to males that reunited, and they did not move more often. The probability of breeding with a new partner was better predicted by female age than by previous breeding success, suggesting that re‐mating was not strongly affected by past breeding performance. Because younger females change breeding sites more frequently than do older females and females that mated with a new partner were younger than females that reunited with their previous mates, we suggest that the tendency of tree swallows to change partners between years is a by‐product of lower site fidelity of younger females rather than a strategy for increasing breeding success.     

Kin Recognition and Incest Avoidance in Toads

Toads, like many amphibians, display breeding site fidelity.Individuals that return to natal ponds to breed are likely toencounter siblings as potential mates. We examined the geneticstructure of Bufo americanus breeding populations at five localitieswithin a 1 km radius. Mitochondrial DNA (mtDNA) haplotypes significantlydiffered among the breeding populations, but haplotype distributionsdid not differ from year to year within breeding sites. Natalphilopatry may account for these results. Although incest isthus possible, siblings rarely mate. Comparing haplotypes oftoads captured while mating, we found only 2 of 86 pairs thatcould possibly be siblings. We suggest that toads might recognizeand actively avoid close kin as mates. Advertisement vocalizationsgiven by males serve as potential cues by which females mightrecognize their kin. We recorded calls of males, in the field,and later obtained rank estimates of their relationships toone another by DNA fingerprinting of the nuclear genome. Theresemblance of males' calls was positively correlated with thesimilarity of their fingerprints. Significant differences incall characteristics were also found among breeding populations.Temporal parameters, rather than frequency components, encodekinship information. Recently developed techniques in moleculargenetics facilitate studies of the heritability of behavioraltraits in natural populations.     

FACTORS AFFECTING THE AGE OF FIRST BREEDING OF THE KITTIWAKE RISSA TRIDACTYLA

At a Kittiwake colony in Northumberland, 80% of those birds which returned to their natal colony to breed were males and these supplied 52% of all male recruits. More females breed away from their natal colony than males. There was no differences in the proportions of young fledged from sites in the centre or at the edge of the colony, or by parents of different experience, which returned to breed. Kittiwakes breed for the first time at ages from 3 to 8 years, but most at 4 or S years old. Males arrive back at the colony at an earlier age than females and breed for the first time one year earlier. Males obtaining sites at the centre of the colony first breed at an earlier age than those at the edges. Neither the age nor the area of first breeding appear to be transmitted from parent to offspring. Males breeding first aged 4 years or younger produced more young than those which first bred aged 5 years or older, despite their partners laying smaller clutches. This difference was most marked among those males recruited to sites in the centre of the colony. The advantage of this earlier breeding is counteracted by a lower survival rate among those males which start to breed at the younger ages. In all breeding Kittiwakes, annual reproductive output increases with experience while annual survival rates decrease. Once they had started to breed, many birds failed to breed in one subsequent season. Nearly 60% of these cases of intermittent breeding occurred in the year following first breeding. Intermittent breeding was most frequent among young birds and among females. It is suggested that each breeding involves a cost to the individual in terms of reduced survival, and that deferred and intermittent breeding are means of guarding survival. A model is proposed whereby the age at which a bird starts to breed, the nesting site which it obtains, and its subsequent breeding strategy result in each individual producing an optimal number of reproducing offspring in its lifetime, relative to its quality.     

How partnerships end in guillemots Uria aalge: chance events, adaptive change, or forced divorce?

Divorce in socially monogamous species can result from differentmechanisms, for example, chance events, active desertion ofthe partner, or the intrusion of a third individual oustingthe partner. We compared the predictions associated with suchmechanisms with data from common guillemots (Uria aalge) breedingon the Isle of May, Scotland. The data cover the years 1982–2005and show a yearly divorce rate of 10.2%. In most divorces (86%),one of the original partners moved to another breeding site,whereas the other bird stayed and bred with a new partner. Onaverage, movers had a significantly lower breeding success afterdivorce, stayers were largely unaffected, whereas the incomingbirds benefited significantly from the change. This patternfits best the predictions of the "forced-divorce" hypothesis,suggesting that many divorces were caused by incoming birdsrather than the original partners or chance events. Althoughwe are unable to document the precise behavioral sequence thatled to divorces, our interpretation is supported by observationsof frequent fights over breeding-site ownership. Our data alsoindicate within-population diversity of divorce mechanisms:some divorces were apparently accidental, others desertion ofpartners and sites if the latter were of low quality. Our studyfinally illustrates that a negative correlation between breedingsuccess and probability of divorce (which our data show) neednot indicate the adaptiveness of divorce for the original partners.Because such a connection has often been made, adaptive divorcemay in general be less common than usually assumed.     

The pair bond and divorce among Oystercatchers Haematopus ostralegus on Skokholm Island, Wales

Mate changes of Oystercatchers were followed over a 15-year period. In 8.0% of instances, the pair split up between breeding seasons even when both members bred the second season. The likelihood of divorce was highest for birds aged five years, the modal age of first breeding. After divorce birds did not select an older or younger mate when re-pairing. Low hatching success increased the chances of divorce. Divorce was not followed by significant changes in clutch-size, laying date or nesting success (although females tended to be more successful).     

Survival of the Orange-tufted Sunbird,Nectarinia osea

Cape Sugarbirds are southern African endemics, found in fynbos vegetation of the Western and Eastern Cape Provinces of South Africa, and they feed almost exclusively on the nectar of Protea inflorescences. Breeding male Sugarbirds are highly territorial and defend resources for themselves, their mates and their offspring. Sugarbirds, however, often leave their breeding territories in search of food during the dry season and return to the same breeding sites each year, thus experiencing an annual cycle of movement from one food source to another. Male territory size as well as breeding success, were determined over two breeding seasons for a population in the Helderberg Nature Reserve in the Western Cape Province. Sugarbird fledgling success was significantly greater for males defending large territories. Males moved or increased the size of their territories between seasons, possibly to improve breeding success, but no changes in territory size were observed during the breeding season.     

The effect of mate change and new partner's age on reproductive success in the barnacle goose, Branta leucopsis

Mate retention frequencies and correlations between mate changeand reproductive performance were estimated in a populationof barnacle geese, Branta leucopsis, breeding on the islandof Gotland in the Baltic. About 90% of the pairs remained togetherfrom one breeding season to the next. Only 2.4% of the pairsdivorced, most mate changes being consequences of the deathof one partner. Divorces were not forecast by low reproductivesuccess, and seemed to be accidental. In the season before matechange, there was no difference in reproductive performance,measured as clutch size, hatching date, and number of fledgedyoung, between faithful pairs and pairs where one partner wassubsequently changed. However, in the first season with a newpartner, clutch size and number of fledged young decreased onaverage. Hence, because mate change led to a reduction in reproductivesuccess, it was concluded that mate retention is advantageous.Our results suggest that this reduction is more likely due tothe lower average age or breeding experience of new partnersthan to the benefits of breeding experience with one particularpartner. [Behav Ecol 1991 ;2:116–122]     

Burrow and mate fidelity in the Little Penguin Eudyptula minor at Lion Island, New South Wales, Australia

Although both mate and nest fidelity are thought to exist in Little Penguins Eudyptula minor , few studies have statistically assessed the relationship between both types of fidelity and breeding success. Observations of the breeding biology of the Little Penguin were carried out between 1990 and 1998 at Lion Island, New South Wales, Australia. The existence of fidelity and the effect of breeding success on subsequent fidelity were assessed for this population. Strong evidence for the existence of nest and mate fidelity was found with high rates of return to the same burrow and mate being recorded for both sexes. For all years of the study, the average number of female and male birds faithful to their nest was 76 and 79%, respectively, with only two noted cases of intraseasonal burrow swapping. The rate of divorce varied widely between years, with a range of 0–40%. A marginal association of a log linear model showed a highly significant relationship ( P  < 0.001) between success of a clutch in one year and fidelity to a mate in the following year. Consequently, parents successfully raising a fledgling in one year were more likely to reunite in the following year than were pairs that attempted unsuccessfully to breed. Pairs failing to fledge chicks were much more likely to separate owing to the absence of one parent in the following year. We were unable to detect any association between breeding success in one year and subsequent change of nest in the next.     

Fitness consequences of divorce in the oystercatcher, Haematopus ostralegus

We investigated the fitness consequences of divorce in oystercatchers. We made a distinction between two types of divorce: in desertions the disruption of the pair bond is initiated by one of the pair members, and in usurpations by a conspecific individual. Survival and reproduction prospects for oystercatchers are largely determined by their social status (nonbreeder or breeding bird at a site of a specific quality). Changes in social status in relation to divorce showed that birds taking the initiative to leave their mate increased in fitness, relative to birds that were forced to leave their partner. Status of individuals that remained in their territory after divorce was unaffected if their mate was expelled, but declined if their mate deserted. Survival after divorce was significantly lower for birds that were expelled than for those deserting. Divorce rate, and especially desertion rate, was higher among occupants of low- than high-quality territories. In general, divorce rate increased following elevated mortality. In high-quality territories usurpations increased with increasing breeder mortality, but at low-quality territories this relation was absent. Desertion rates were similarly related to mortality in both territory types. Divorce participants thus differed strongly in their fitness prospects, depending on the type of divorce, the role played in the divorce and the quality of the territory where divorce took place. Studies that do not observe the birds during divorce cannot determine the type of divorce and the role played by the individuals, and this may lead to misleading conclusions on the costs and benefits of divorce. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Mate fidelity in monogamous birds: a re-examination of the Procellariiformes

Mate retention is classically considered advantageous for reproduction in monogamous birds: because of their low fecundity, long-lived species should show the highest year-to-year mate fidelity. However, this hypothesis remains controversial: several studies have found no correlation between mate fidelity and longevity, possibly because they did not control for potential confounding factors on each of these parameters, and one study found a negative correlation in the Procellariiformes (albatrosses and petrels). We re-examined the relations between mate fidelity and longevity, and between mate fidelity and site fidelity, in this group, using our data on 13 species and data from the literature, and after eliminating confounding factors. Procellariiformes are the most long lived of birds despite important interspecific variation in body size, and they show strong mate fidelity and bear high costs of divorce. All species lay only one egg, and the most long lived breed biennially. Because large organisms live longer than small ones and their reproductive effort is lower, we had to control for breeding frequency and body size. Mate fidelity and adult life expectancy were positively correlated, regardless of whether we controlled for these two parameters. We also evaluated whether mate fidelity was related to site fidelity. Biennial albatrosses show high mate fidelity, but low nest fidelity, although they are extremely faithful to a small area around their previous nest. After controlling for body size, adult life expectancy and breeding frequency, we found no correlation between mate fidelity and site fidelity. Because divorce is costly and mate retention advantageous in Procellariiformes, we suggest that mate fidelity does not passively result from site fidelity in these species. Rather, site fidelity would be a means for pairs to reunite, with sites serving as meeting points. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.