Hormone levels predict individual differences in reproductive success in a passerine bird

Hormones mediate major physiological and behavioural components of the reproductive phenotype of individuals. To understand basic evolutionary processes in the hormonal regulation of reproductive traits, we need to know whether, and during which reproductive phases, individual variation in hormone concentrations relates to fitness in natural populations. We related circulating concentrations of prolactin and corticosterone to parental behaviour and reproductive success during both the pre-breeding and the chick-rearing stages in both individuals of pairs of free-living house sparrows, Passer domesticus. Prolactin and baseline corticosterone concentrations in pre-breeding females, and prolactin concentrations in pre-breeding males, predicted total number of fledglings. When the strong effect of lay date on total fledgling number was corrected for, only pre-breeding baseline corticosterone, but not prolactin, was negatively correlated with the reproductive success of females. During the breeding season, nestling provisioning rates of both sexes were negatively correlated with stress-induced corticosterone levels. Lastly, individuals of both sexes with low baseline corticosterone before and high baseline corticosterone during breeding raised the most offspring, suggesting that either the plasticity of this trait contributes to reproductive success or that high parental effort leads to increased hormone concentrations. Thus hormone concentrations both before and during breeding, as well as their seasonal dynamics, predict reproductive success, suggesting that individual variation in absolute concentrations and in plasticity is functionally significant, and, if heritable, may be a target of selection.     

THE FOUNDING OF A NEW POPULATION OF DARWIN'S FINCHES

We report the natural colonization of the small Galápagos island Daphne Major by the large ground finch (Geospiza magnirostris). Immigrants of this species were present in every year of a 22-yr study, 1973–1994. Typically they arrived after a breeding season and left at the beginning of the next one. Geospiza magnirostris bred on the island for the first time in the exceptionally wet El Niño year of 1982–1983, and bred in all subsequent years except drought years. In agreement with theoretical expectations the frequency of inbreeding was unusually high. Pronounced fluctuating asymmetry in tarsus length, together with slightly reduced breeding success of inbreeding pairs, suggests a low level of inbreeding depression. Despite this, the population increased from 5 breeding individuals in 1983 to 20 breeding individuals in 1992, and probably more than twice that number in 1993, largely through recruitment of locally born birds.     

Individual fitness and the effects of a changing climate on the cessation and length of the breeding period using a 34‐year study of a temperate songbird

Studies of the phenological responses of animals to climate change typically emphasize the initiation of breeding although climatic effects on the cessation and length of the breeding period may be as or more influential of fitness. We quantified links between climate, the cessation and length of the breeding period, and individual survival and reproduction using a 34‐year study of a resident song sparrow (Melospiza melodia) population subject to dramatic variation in climate. We show that the cessation and length of the breeding period varied strongly across years, and predicted female annual fecundity but not survival. Breeding period length was more influential of fecundity than initiation or cessation of breeding alone. Warmer annual temperature and drier winters and summers predicted an earlier cessation of breeding. Population density, the date breeding was initiated, a female's history of breeding success, and the number of breeding attempts initiated previously also predicted the cessation of breeding annually, indicating that climatic, population, and individual factors may interact to affect breeding phenology. Linking climate projections to our model results suggests that females will both initiate and cease breeding earlier in the future; this will have opposite effects on individual reproductive rate because breeding earlier is expected to increase fecundity, whereas ceasing breeding earlier should reduce it. Identifying factors affecting the cessation and length of the breeding period in multiparous species may be essential to predicting individual fitness and population demography. Given a rich history of studies on the initiation of breeding in free‐living species, re‐visiting those data to estimate climatic effects on the cessation and length of breeding should improve our ability to predict the impacts of climate change on multiparous species.     

Estimating fluctuating selection in age-structured populations

In age-structured populations, viability and fecundity selection of varying strength may occur in different age classes. On the basis of an original idea by Fisher of weighting individuals by their reproductive value, we show that the combined effect of selection on traits at different ages acts through the individual reproductive value defined as the stochastic contribution of an individual to the total reproductive value of the population the following year. The selection differential is a weighted sum of age-specific differentials that are the covariances between the phenotype and the age-specific relative fitness defined by the individual reproductive value. This enables estimation of weak selection on a multivariate quantitative character in populations with no density regulation by combinations of age-specific linear regressions of individual reproductive values on the traits. Demographic stochasticity produces random variation in fitness components in finite samples of individuals and affects the statistical inference of the temporal average directional selection as well as the magnitude of fluctuating selection. Uncertainties in parameter estimates and test power depend strongly on the demographic stochasticity. Large demographic variance results in large uncertainties in yearly estimates of selection that complicates detection of significant fluctuating selection. The method is illustrated by an analysis of age-specific selection in house sparrows on a fitness-related two-dimensional morphological trait, tarsus length and body mass of fledglings.     

Improvement of reproductive performance with age and breeding experience depends on recruitment age in a long‐lived seabird

Reproductive success increases with age in many species, however, the underlying mechanisms remain unclear and several hypotheses have been proposed to explain age‐related improvements in reproductive output. In this contribution we investigated the effects of age, recruitment age, breeding experience and sex on reproductive performance during the early breeding career in the common tern Sterna hirundo using long‐term individual‐based data. We used measurements of performance, which spanned the entire breeding process: clutch size, hatching success, fledging success and fledglings per pair. Longitudinal analyses within individuals showed a clear increase with age in all performance measures. Furthermore, a significant change in reproductive performance was found between first time‐ and experienced breeders. Recruitment age had a strong influence on hatching and fledging success: two‐year‐old recruits had significantly lower reproductive success than birds which recruited at older ages, but the increase in breeding performance with experience was stronger in young recruits. Comparing age and experience effects, age effects were more pronounced during the first breeding attempts, whereas experience effects were also visible in subsequent breeding attempts. The degree of intra‐individual improvements in reproductive performance is due to a complex interplay of age at first breeding and experience. The results strongly support the constraint hypothesis.     

The quality and the timing hypotheses evaluated using data on great reed warblers

The seasonal decline in reproductive success observed in many animal species may be caused by timing per se (timing hypothesis) or by variation in phenotypic quality between early and late breeding females (quality hypothesis). To distinguish between these two hypotheses, several studies of birds have used clutch removal experiments to manipulate breeding date. However, removal experiments also increase the females' previous reproductive effort due to the production of an extra clutch and a longer incubation period. According to life-history theory an increase in reproductive effort lowers future reproduction. Hence, life-history theory predicts lowered success of replacement broods for other reasons than expected from the timing hypothesis. Female great reed warblers, Acrocephalus arundinaceus , studied in Sweden are frequently exposed to nest predation, after which many lay replacement clutches. In order to examine possible effects of previous reproductive effort on different fitness components, we analysed the re-laying frequency and the reproductive success of replacement broods in relation to time of the season and previous reproductive effort (measured as the length of the previous breeding attempt, LPB). In clutches of re-laying females both the number of fledglings and the proportion of recruits were negatively correlated with LPB, whereas re-laying frequency and clutch size were not related to LPB. We expect such relationships to be present also among other species. Consequently, the use of replacement clutches, as for example in clutch removal experiments, in evaluations of the cause of the often observed seasonal decline in various fitness components, might exaggerate the importance of the timing hypothesis over the quality hypothesis.     

Constraints on, and determinants of, the annual number of breeding attempts in the multi-brooded Black Redstart Phoenicurus ochruros

With reference to models predicting patterns of reproduction in multi-brooded species, I analysed some of the factors potentially affecting the number of breeding attempts per season made by female Black Redstarts Phoenicurus ochruros in the central Swiss Alps. The maximum number of successful broods per female per season was three, the median two. The proportion of females initiating only a single clutch per season varied from 16 to 58% over 10 years. Variation in the frequency of single brooding between years was positively related to the date of the onset of breeding. Egg-laying started after a temperature-related threshold value was crossed in April. Females frequently re-nested before fledglings of the previous brood were independent. Short interbrood intervals were associated with triple brooding. The interbrood interval shortened during the breeding season. Intra-individual variation in the number of breeding attempts per season was partitioned according to female age, laying date of the season's first egg (standardized in relation to the median date of first laying within a year) and the occurrence of breeding failures before the ultimate nesting attempt. However, mortality during the season often terminated breeding early, in particular among novice breeder females. Seasonal reproductive success increased linearly with each additional breeding attempt. The productivity of a breeding attempt was independent of the time in the season. Increased reproductive effort affected neither current or future survival nor reproduction of females. The reproductive patterns and trade-offs in multi-brooded Black Redstarts contrast in many aspects with those found in single-brooded species. They are in accordance with models predicting that multi-brooded species are selected to start breeding as early as possible, continue breeding for as long as conditions are suitable, and save time by overlapping broods.     

Unpredictable food supply modifies costs of reproduction and hampers individual optimization

Investment into the current reproductive attempt is thought to be at the expense of survival and/or future reproduction. Individuals are therefore expected to adjust their decisions to their physiological state and predictable aspects of environmental quality. The main predictions of the individual optimization hypothesis for bird clutch sizes are: (1) an increase in the number of recruits with an increasing number of eggs in natural broods, with no corresponding impairment of parental survival or future reproduction, and (2) a decrease in the fitness of parents in response to both negative and positive brood size manipulation, as a result of a low number of recruits, poor future reproduction of parents, or both. We analysed environmental influences on costs and optimization of reproduction on 6 years of natural and experimentally manipulated broods in a Central European population of the collared flycatcher. Based on dramatic differences in caterpillar availability, we classified breeding seasons as average and rich food years. The categorization was substantiated by the majority of present and future fitness components of adults and offspring. Neither observational nor experimental data supported the individual optimization hypothesis, in contrast to a Scandinavian population of the species. The quality of fledglings deteriorated, and the number of recruits did not increase with natural clutch size. Manipulation revealed significant costs of reproduction to female parents in terms of future reproductive potential. However, the influence of manipulation on recruitment was linear, with no significant polynomial effect. The number of recruits increased with manipulation in rich food years and tended to decrease in average years, so control broods did not recruit more young than manipulated broods in any of the year types. This indicates that females did not optimize their clutch size, and that they generally laid fewer eggs than optimal in rich food years. Mean yearly clutch size did not follow food availability, which suggests that females cannot predict food supply of the brood-rearing period at the beginning of the season. This lack of information on future food conditions seems to prevent them from accurately estimating their optimal clutch size for each season. Our results suggest that individual optimization may not be a general pattern even within a species, and alternative mechanisms are needed to explain clutch size variation.     

Individual differences in developmental precision and fluctuating asymmetry: a model and its implications

In many studies, fluctuating asymmetry (FA) has been used as a measure of individual differences in developmental imprecision. A model of how variation in developmental imprecision is associated with variation in asymmetry is described and applied to important issues about FA. If individual differences in developmental imprecision exist, asymmetry due to developmental error should be leptokurtically distributed. Moreover, the greater the magnitude of individual differences, the greater the leptokurtosis. Asymmetry purportedly due to developmental error in a variety of species is indeed leptokurtically distributed. The level of leptokurtosis suggests that the CV in individual differences in underlying developmental imprecision is generally 20–25, consistent with it being a fitness trait. In addition, data suggest that: (1) the individual differences that underlie the developmental imprecision of different traits are largely shared across traits and not trait-specific; (2) the heritability of these individual differences may average between 35 and 55%, despite small heritabilities of individual trait FAs; and (3) correlations between FA and fitness traits or components suggest high correlations between underlying variation in developmental precision and fitness in many species. Theoretical implications are discussed.     

Does prior breeding experience improve reproductive success in collared flycatcher females?

In order to assess the importance of breeding experience in shaping age-specific patterns of reproduction an experiment on collared flycatchers (Ficedula albicollis) was performed. Experimental females were restrained from reproduction by destroying their nests after prolonged incubation. Unmanipulated females were used as controls. Only young, first time breeders were considered. A year after manipulation, experimental (inexperienced) and control (experienced) females were compared for a number of breeding traits. They did not differ with respect to laying date, clutch size, number of fledglings and number of local recruits. However, fledglings from broods of inexperienced females were significantly lighter and had shorter tarsi. This suggests that prior experience of feeding young may indeed influence the future reproductive performance in altricial bird species such as the collared flycatcher.     

Factors influencing double brooding in Eurasian Hoopoes Upupa epops

Double brooding may be a good strategy for short‐lived species to maximize annual and lifetime reproductive success (ARS and LRS, respectively). Nevertheless, there is typically individual variation in the probability of producing a second clutch. Here we evaluate factors that influence the decision to double brood in the Eurasian Hoopoe Upupa epops. Analyses of an 11‐year dataset showed that 36% of the females and 21% of the males produced a second clutch after successfully raising a first clutch. Double‐brooded females had higher ARS (9.1 ± 1.9 fledglings; mean ± se) and LRS (0.93 ± 0.08 recruits) than single‐brooded females (ARS: 4.5 ± 2.1 fledglings; LRS: 0.36 ± 0.03 recruits). This suggests that double brooding is adaptive in Hoopoes, and raises the question of why most individuals only produce one clutch per season. The probability of double brooding varied only slightly between years, suggesting that it is influenced by individual characteristics rather than by external, population‐level environmental factors. In both sexes, the probability of double brooding increased with earlier timing of the first clutch, and the timing of reproduction was the most important factor influencing reproductive success. The latter is likely to be mediated by changes in resources during the season. The probability of double brooding also increased slightly with female age, due to differences in intrinsic quality among females rather than to a gain in experience. In contrast to many other studies, the probability of double brooding increased with an increasing number of fledglings from the first clutch, suggesting that it is a strategy of individuals of high quality. Taken together, we show that the individual quality of the breeder and the timing of their first clutches are key factors influencing the decision to double brood, and thereby that they are important determinants of reproductive performance in Eurasian Hoopoes.     

QUANTITATIVE GENETICS OF FITNESS TRAITS IN A WILD POPULATION OF PHLOX

To determine if the evolution of fitness traits in the annual plant, Phlox drummondii, is constrained by lack of genetic variation, we calculated the heritability and genetic correlation of 16 traits in a field population. Full- and half-sib families of seeds were generated in the greenhouse and planted back into the study population. Of 855 seeds that germinated, 609 survived to produce fruit. For each plant we measured several aspects of plant size and three components of female fecundity: total number of fruits produced, number of seeds per fruit, and mass of individual seeds. Heritability of traits ranged from 0.00 to 0.15. Several traits showed significant levels of additive genetic variance, but we found no evidence of additive genetic variance in components of female fecundity and no evidence of negative genetic correlation between fitness traits. These results suggest that evolution in this population would be constrained by lack of heritable variation in fitness traits.     

Multiple breeding in the Great Tit. A trade-off between successive reproductive attempts?

1. Multiple breeding (raising more than one batch of young per breeding season) is a common life-history tactic that has received very little attention. A simple static optimization model was developed, applicable to iteroparous animals with parental care, that predicts: (1) when an animal should be a multiple breeder, (2) the optimal interval between successive clutches, and (3) when a clutch should be deserted.
2. More specifically, it was predicted that (I) animals should invest less in a clutch when it is followed by another clutch in the same season, and (II) as a consequence the contribution of the first clutch to the parents' fitness will be reduced.
3. These predictions were tested experimentally in a population of Great Tits ( ParusM. major L.). The experiment consisted of the removal of second clutches. When second clutches were removed, parents tended their first brood fledglings for a longer period than undisturbed control pairs, in agreement with the first prediction. Removal of second clutches did not affect fledgling survival, but recruits enjoyed higher reproductive success in their first year of breeding when the second clutch of their parents had been removed, in agreement with the second prediction.
4. It is concluded that a trade-off exists between successive reproductive attempts, which will affect the optimal rate of reproductive attempts.     

The significance of subadult plumage in Darwin's finches, Geospiza fortis

I studied delayed plumage maturation, a common phenomenon inpasserine birds, in a species of Darwin's finch, Geospiza fortis,on Isla Daphne Major, Galapagos, Ecuador. To address the questionof why males advertise their youth and immaturity with subadultplumage, I determined the fitness consequences of possessingdifferent plumage states at different ages. Rate of entry intoadult plumage is both variable and heritable. In two cohorts,individuals that took longer to enter full adult plumage surviveddry conditions significantly better than those that acquiredadult plumage quickly. Behavioral observations helped revealthe reasons for this higher survival. Advantages arose fromthe avoidance of conflict that occurred by signaling subordinatestatus with subadult plumage in both the nonbreeding and breedingseasons. Although blacker males had a significantly lower survival,those black males that did survive had a reproductive advantagein their first breeding season over males with less black intheir plumage from the same cohort and raised significantlymore offspring to fledging. Thus the study demonstrates selectionon the length of the subadult plumage stage. [Behav Ecol 1990;l:161–170]     

PHENOTYPIC AND GENETIC EFFECTS OF HYBRIDIZATION IN DARWIN'S FINCHES

Morphological consequences of hybridization were studied in a group of three interbreeding species of Darwin's finches on the small Galápagos island of Daphne Major in the inclusive years 1976 to 1992. Geospiza fortis bred with G. scandens and G. fuliginosa. Although interbreeding was always rare (< 5%), sufficient samples of measurements of hybrids and backcrosses were accumulated for analysis. Five beak and body dimensions and mass were measured, and from these two synthetic (principal-component) traits were constructed. All traits were heritable in two of the interbreeding species (G. fuliginosa were too rare to be analyzed) and in the combined samples of F, hybrids and backcrosses to G. fortis. In agreement with expectations from a model of polygenic inheritance, hybrid and backcross classes were generally phenotypically intermediate between the breeding groups that had produced them. Hybridization increased additive genetic and environmental variances, increased heritabilities to a moderate extent, and generally strengthened phenotypic and genetic correlations. New additive genetic variance introduced by hybridization is estimated to be two to three orders of magnitude greater than that introduced by mutation. Enhanced variation facilitates directional evolutionary change, subject to constraints arising from genetic correlations between characters. The Darwin's finch data suggest that these constraints become stronger when species with similar proportions hybridize, but some become weaker when the interbreeding species have different allometries. This latter effect of hybridization, together with an enhancement of genetic variation, facilitates evolutionary change in a new direction.     

Temperature fluctuations during development reduce male fitness and may limit adaptive potential in tropical rainforest Drosophila

Understanding the potential for organisms to tolerate thermal stress through physiological or evolutionary responses is crucial given rapid climate change. Although climate models predict increases in both temperature mean and variance, such tolerances are typically assessed under constant conditions. We tested the effects of temperature variability during development on male fitness in the rainforest fly Drosophila birchii, by simulating thermal variation typical of the warm and cool margins of its elevational distribution, and estimated heritabilities and genetic correlations of fitness traits. Reproductive success was reduced for males reared in warm (mean 24 °C) fluctuating (±3 °C) vs. constant conditions but not in cool fluctuating conditions (mean 17 °C), although fluctuations reduced body size at both temperatures. Male reproductive success under warm fluctuating conditions was similar to that at constant 27 °C, indicating that briefly exceeding critical thermal limits has similar fitness costs to continuously stressful conditions. There was substantial heritable variation in all traits. However, reproductive success traits showed no genetic correlation between treatments reflecting temperature variation at elevational extremes, which may constrain evolutionary responses at these ecological margins. Our data suggest that even small increases in temperature variability will threaten tropical ectotherms living close to their upper thermal limits, both through direct effects on fitness and by limiting their adaptive potential.     

Evaluating Habitat Quality for the Dusky Flycatcher

Abstract: We studied the dusky flycatcher (Empidonax oberholseri) at 8 sites in central Idaho, USA, in 2002 and 2003 to examine relationships among vegetation cover, density of breeding conspecifics, and indicators of habitat quality. Number of breeding territories and number of fledglings per hectare were positively associated, suggesting that the dusky flycatcher experienced increased reproductive success where it bred at the highest densities. However, the relationships between nesting success, annual reproductive success, number of fledglings per hectare, and amount of understory cover showed substantial annual variation. Nesting success did not differ significantly across sites or between years. Both reproductive success, expressed as young produced per hectare (range: 0.34–3.09 in 2002 and 0.79–3.82 in 2003) and young produced per nesting attempt (range: 0.71–2.78 in 2002 and 1.11–3.10 in 2003), differed across study sites in each year. Mean clutch size did not differ significantly among sites or years. Mean egg weight showed significant variation across some sites within years and was associated negatively with the 3 measures of reproductive success in 2002, although small sample sizes prevented reliable inference about the appropriateness of this measure as an indicator of habitat quality. Mean clutch size and mean egg weight were not associated with vegetation cover variables. Thus, dusky flycatcher reproductive success showed inconsistencies with individual vegetation measurements at the site scale. Forest managers who use vegetation treatments to increase amounts of understory shrub cover (e.g., by removing portions of the overstory conifer canopy) should increase densities of this species and, in turn, increase number of fledglings produced, but these responses appear to be better measured at the territory or nest scales than at the stand or site scales.     

Environmental conditions affect the magnitude of inbreeding depression in survival of Darwin's finches

Understanding the fitness consequences of inbreeding (inbreeding depression) is of importance to evolutionary and conservation biology. There is ample evidence for inbreeding depression in captivity, and data from wild populations are accumulating. However, we still lack a good quantitative understanding of inbreeding depression and what influences its magnitude in natural populations. Specifically, the relationship between the magnitude of inbreeding depression and environmental severity is unclear. We quantified inbreeding depression in survival and reproduction in populations of cactus finches (Geospiza scandens) and medium ground finches (Geospiza fortis) living on Isla Daphne Major in the Galápagos Archipelago. Our analyses showed that inbreeding strongly reduced the recruitment probability (probability of breeding given that an adult is alive) in both species. Additionally, in G. scandens, first-year survival of an offspring with f = 0.25 was reduced by 21% and adults with f = 0.25 experienced a 45% reduction in their annual probability of survival. The magnitude of inbreeding depression in both adult and juvenile survival of this species was strongly modified by two environmental conditions, food availability and number of competitors. In juveniles, inbreeding depression was only present in years with low food availability, and in adults inbreeding depression was five times more severe in years with low food availability and large population sizes. The combination of relatively severe inbreeding depression in survival and the reduced recruitment probability led to the fact that very few inbred G. scandens ever succeeded in breeding. Other than recruitment probability, no other trait showed evidence of inbreeding depression in G. fortis, probably for two reasons: a relatively high rate of extrapair paternity (20%), which may lead to an underestimate of the apparent inbreeding depression, and low sample sizes of highly inbred G. fortis, which leads to low statistical power. Using data from juvenile survival, we estimated the number of lethal equivalents carried by G. scandens, G. fortis, and another congener, G. magnirostris. These results suggest that substantial inbreeding depression can exist in insular populations of birds, and that the magnitude of the inbreeding depression is a function of environmental conditions.     

Do Zebra Finch Parents Fail to Recognise Their Own Offspring?

Individual recognition systems require the sender to be individually distinctive and the receiver to be able to perceive differences between individuals and react accordingly. Many studies have demonstrated that acoustic signals of almost any species contain individualized information. However, fewer studies have tested experimentally if those signals are used for individual recognition by potential receivers. While laboratory studies using zebra finches have shown that fledglings recognize their parents by their "distance call", mutual recognition using the same call type has not been demonstrated yet. In a laboratory study with zebra finches, we first quantified between-individual acoustic variation in distance calls of fledglings. In a second step, we tested recognition of fledgling calls by parents using playback experiments. With a discriminant function analysis, we show that individuals are highly distinctive and most measured parameters show very high potential to encode for individuality. The response pattern of zebra finch parents shows that they do react to calls of fledglings, however they do not distinguish between own and unfamiliar offspring, despite individual distinctiveness. This finding is interesting in light of the observation of a high percentage of misdirected feedings in our communal breeding aviaries. Our results demonstrate the importance of adopting a receiver's perspective and suggest that variation in fledgling contact calls might not be used in individual recognition of offspring.     

Direct fitness of group living mammals varies with breeding strategy, climate and fitness estimates

1. Understanding how variation in fitness relates to variation in group living remains critical to determine whether this major aspect of social behaviour is currently adaptive. 2. Available evidence in social mammals aimed to examine this issue remains controversial. Studies show positive (i.e. potentially adaptive), neutral or even negative fitness effects of group living. 3. Attempts to explain this variation rely on intrinsic and extrinsic factors to social groups. Thus, relatively more positive fitness effects are predicted in singularly breeding as opposed to plural breeding species. Fitness effects of sociality in turn may depend on ecological conditions (i.e. extrinsic factors) that influence associated benefits and costs. 4. We used meta-analytic tools to review how breeding strategy or ecological conditions influence the effect size associated with direct fitness-sociality relationships reported in the mammalian literature. Additionally, we determined how taxonomic affiliation of species studied, different fitness and sociality measures used, and major climatic conditions of study sites explained any variation in direct fitness effect size. 5. We found group living had modest, yet positive effects on direct fitness. This generally adaptive scenario was contingent not only upon breeding strategy and climate of study sites, but also on fitness measures examined. Thus, positive and significant effects characterized singular as opposed to plural breeding strategies. 6. We found more positive fitness effects on studies conducted in tropical as opposed to temperate or arid climates. More positive and significant effects were noted on studies that relied on group fecundity, male fecundity and offspring survival as measures of fitness. 7. To conclude, direct fitness consequences of mammalian group living are driven by interspecific differences in breeding strategy and climate conditions. Other factors not examined in this study, namely individual variation in direct and indirect fitness benefits and potential interactions between social and ecological conditions, may be important and require further studies.