Variation In Reproductive Success Research Articles

BET HEDGING IN A GUILD OF DESERT ANNUALS

Evolutionary bet hedging encapsulates the counterintuitive idea that organisms evolve traits that reduce short-term reproductive success in favor of longer-term risk reduction. It has been widely investigated theoretically, and many putative examples have been cited including practical ones such as the dormancy involved in microbe and weed persistence. However, long-term data on demographic variation from the actual evolutionarily relevant environments have been unavailable to test for its mechanistic relationship to alleged bet hedging traits. I report an association between delayed germination (a bet hedging trait) and risk using a 22-year data set on demographic variation for 10 species of desert annual plants. Species with greater variation in reproductive success (per capita survival from germination to reproduction x per capita fecundity of survivors) were found to have lower average germination fractions. This provides a definitive test using realistic data on demographic variance that confirms the life history prediction for bet hedging. I also showed that the species with greater long-term demographic variation tended to be the ones with greater sensitivity of reproductive success to variation among years in growing-season precipitation.

ITEROPARITY IN THE VARIABLE ENVIRONMENT OF THE SALAMANDERAMBYSTOMA TIGRINUM

Simultaneous estimation of survival, reproduction, and movement is essential to understanding how species maximize lifetime reproduction in environments that vary across space and time. We conducted a four-year, capture-recapture study of three populations of eastern tiger salamanders (Ambystoma tigrinum tigrinum) and used multistate mark-recapture statistical methods to estimate the manner in which movement, survival, and breeding probabilities vary under different environmental conditions across years and among populations and habitats. We inferred how individuals may mitigate risks of mortality and reproductive failure by deferring breeding or by moving among populations. Movement probabilities among populations were extremely low despite high spatiotemporal variation in reproductive success and survival, suggesting possible costs to movements among breeding ponds. Breeding probabilities varied between wet and dry years and according to whether or not breeding was attempted in the previous year. Estimates of survival in the nonbreeding, forest habitat varied among populations but were consistent across time. Survival in breeding ponds was generally high in years with average or high precipitation, except for males in an especially ephemeral pond. A drought year incurred severe survival costs in all ponds to animals that attempted breeding. Female salamanders appear to defer these episodic survival costs of breeding by choosing not to breed in years when the risk of adult mortality is high. Using stochastic simulations of survival and breeding under historical climate conditions, we found that an interaction between breeding probabilities and mortality limits the probability of multiple breeding attempts differently between the sexes and among populations.

Reduced X-Linked Diversity in Derived Populations of House Mice

Contrasting patterns of X-linked vs. autosomal diversity may be indicative of the mode of selection operating in natural populations. A number of observations have shown reduced X-linked (or Z-linked) diversity relative to autosomal diversity in various organisms, suggesting a large impact of genetic hitchhiking. However, the relative contribution of other forces such as population bottlenecks, variation in reproductive success of the two sexes, and differential introgression remains unclear. Here, we survey 13 loci, 6 X-linked and 7 autosomal, in natural populations of the house mouse (Mus musculus) subspecies complex. We studied seven populations of three different subspecies, the eastern house mouse M. musculus castaneus, the central house mouse M. m. musculus, and the western house mouse M. m. domesticus, including putatively ancestral and derived populations for each. All populations display lower diversity on the X chromosomes relative to autosomes, and this effect is most pronounced in derived populations. To assess the role of demography, we fit the demographic parameters that gave the highest likelihood of the data using coalescent simulations. We find that the reduction in X-linked diversity is too large to be explained by a simple demographic model in at least two of four derived populations. These observations are also not likely to be explained by differences in reproductive success between males and females. They are consistent with a greater impact of positive selection on the X chromosome, and this is supported by the observation of an elevated K(A) and elevated K(A)/K(S) ratios on the rodent X chromosome. A second contribution may be that the X chromosome less readily introgresses across subspecies boundaries.

Maternal Investment of the Virunga Mountain Gorillas

AbstractThe Trivers & Willard hypothesis (TWH) predicts that females with more resources should bias their maternal investment toward offspring of the sex that is most likely to benefit from those additional resources. This paper examines the sex allocation of 61 female mountain gorillas (Gorilla beringei beringei) of the Virunga volcanoes, Rwanda from 1967 to 2004. Like most highly dimorphic, polygynous mammals, mountain gorillas are expected to show greater variance in reproductive success among males than females, so mothers in good condition should bias their investment toward sons. Using dominance rank as the indicator of maternal condition, the TWH was tentatively supported by our results with interbirth intervals (IBI). Dominant mothers had longer IBI following the birth of sons, relative to the longer IBI that subordinate mothers had with daughters. In contrast, maternal condition did not have a significant effect on birth sex ratios. We also found no significant relationships with other variables that might influence birth sex ratios (e.g., maternal age, parity, or group size), and the overall birth sex ratio was not significantly different from a 50:50 split. Collectively, our results suggest that female mountain gorillas do not control the sex ratio of their offspring at birth, but they may adjust their subsequent maternal investment. This conclusion is consistent with recurring questions about whether any adjustments in birth sex ratios occur in primates.

Effective population size of steelhead trout: influence of variance in reproductive success, hatchery programs, and genetic compensation between life‐history forms

The effective population size is influenced by many biological factors in natural populations. To evaluate their relative importance, we estimated the effective number of breeders per year (Nb) and effective population size per generation (Ne) in anadromous steelhead trout (Oncorhynchus mykiss) in the Hood River, Oregon (USA). Using demographic data and genetic parentage analysis on an almost complete sample of all adults that returned to the river over 15 years (>15,000 individuals), we estimated Nb for 13 run years and Ne for three entire generations. The results are as follows: (i) the ratio of Ne to the estimated census population size (N) was 0.17-0.40, with large variance in reproductive success among individuals being the primary cause of the reduction in Ne/N; (ii) fish from a traditional hatchery program (Htrad: nonlocal, multiple generations in a hatchery) had negative effects on Nb, not only by reducing mean reproductive success but also by increasing variance in reproductive success among breeding parents, whereas no sign of such effects was found in fish from supplementation hatchery programs (Hsupp: local, single generation in a hatchery); and (iii) Nb was relatively stable among run years, despite the widely fluctuating annual run sizes of anadromous adults. We found high levels of reproductive contribution of nonanadromous parents to anadromous offspring when anadromous run size is small, suggesting a genetic compensation between life-history forms (anadromous and nonanadromous). This is the first study showing that reproductive interaction between different life-history forms can buffer the genetic impact of fluctuating census size on Ne.

Maternal androgens in the pied flycatcher: timing of breeding and within-female consistency

Maternal hormones can have substantial phenotypic effects in the progeny of many vertebrates. It has been proposed that mothers adaptively adjust hormone levels experienced by particular young to optimize their reproductive output. In birds, systematic variation in egg hormone levels has been related to different female reproductive strategies. Because in many bird species prospects of the offspring change seasonally and with brood number, strategic adjustment of yolk androgen levels would be expected. To test this idea, we induced pied flycatcher (Ficedula hypoleuca) females to nest twice during the same season by removing their first clutches shortly after clutch completion. We collected eggs of first and replacement clutches to measure yolk concentrations of androstenedione (A4) and testosterone (T) and captured the females that laid these clutches for phenotypic measurements. Although average egg androgen levels were remarkably consistent within females, hormone patterns differed considerably between first and replacement clutches. Eggs of replacement clutches were heavier with larger yolks compared to first clutches, but they contained on average lower levels of androgens. Within clutches, androgen concentration increased over the laying sequence in the first clutch, but decreased or remained more constant over the laying sequence in the replacement clutch. Mean yolk T, but not A4 levels, were negatively associated with laying date for both breeding attempts. Moreover, females in good body condition produced eggs containing lower levels of androgens than females in poor condition. Our results are consistent with the idea that differences in yolk androgen levels may be one mechanism underlying seasonal variation in reproductive success and it is possible that changes in egg androgen patterns may reflect a change in female reproductive strategy. High within-female consistency also highlights the possibility that there may be some underlying genetic variation in yolk androgen levels.

Giant Galápagos tortoises; molecular genetic analyses identify a trans-island hybrid in a repatriation program of an endangered taxon

BackgroundGiant Galápagos tortoises on the island of Española have been the focus of an intensive captive breeding-repatriation programme for over 35 years that saved the taxon from extinction. However, analysis of 118 samples from released individuals indicated that the bias sex ratio and large variance in reproductive success among the 15 breeders has severely reduced the effective population size (Ne).ResultsWe report here that an analysis of an additional 473 captive-bred tortoises released back to the island reveals an individual (E1465) that exhibits nuclear microsatellite alleles not found in any of the 15 breeders. Statistical analyses incorporating genotypes of 304 field-sampled individuals from all populations on the major islands indicate that E1465 is most probably a hybrid between an Española female tortoise and a male from the island of Pinzón, likely present on Española due to human transport.ConclusionRemoval of E1465 as well as its father and possible (half-)siblings is warranted to prevent further contamination within this taxon of particular conservation significance. Despite this detected single contamination, it is highly noteworthy to emphasize the success of this repatriation program conducted over nearly 40 years and involving release of over 2000 captive-bred tortoises that now reproduce in situ. The incorporation of molecular genetic analysis of the program is providing guidance that will aid in monitoring the genetic integrity of this ambitious effort to restore a unique linage of a spectacular animal.

FEMALE-BIASED SEX RATIO ARISES AFTER PARENTAL CARE IN THE SEXUALLY DIMORPHIC WAVED ALBATROSS (PHOEBASTRIA IRRORATA)

Abstract In response to evidence of sexual segregation at foraging grounds as well as male-biased band recoveries, we investigated the ontogeny of the female- biased adult sex ratio in the Waved Albatross (Phoebastria irrorata), an IUCN “critically endangered species” essentially endemic to Isla Espanola, Galapagos, Ecuador. Using a molecular technique to determine the sex of chicks and adults and known fate analysis of chicks during rearing, we found no evidence of a sex-ratio bias at hatching or fledging in three consecutive years with variable reproductive success. Although male chicks were significantly larger than females at fledging, survival to fledging of a large sample of male and female chicks did not differ. The sex ratio among a cohort of young adults at approximately the age of first breeding (eight years) also did not differ significantly from parity. Differential adult mortality, including male-biased mortality in fisheries, is the most probable cause of a female- biased population sex rati...

Core area quality is associated with variance in reproductive success among female chimpanzees at Kibale National Park

Female East African chimpanzees, Pan troglodytes schweinfurthii, tend to range apart from each other in dispersed core areas, and they have dominance interactions with each other so rarely that it is difficult for observers to assess a dominance hierarchy. Nevertheless female chimpanzees can have high variance in fitness. Here, we test the hypothesis that female chimpanzee fitness variance is associated with variation in the foraging quality of their ranges. We studied range usage of 21 wild adult female chimpanzees within the Kanyawara community, Kibale National Park, Uganda. Core areas of individuals remained stable over a 9-year period and varied in their density of preferred foods. Females in neighbourhoods containing more preferred foods had elevated ovarian hormone production, shorter birth intervals and higher infant survivorship. Our results thus suggest that superior access to food may have enabled some community females to reproduce more successfully than others. Although dominance interactions are less frequent among females than among males of this species, we propose that the intensity of selection on intrasexual competition may be similar between the sexes. We discuss potential applications to other fission–fusion species.

Benefits of large broods by higher chick survival and better territories in a precocial shorebird

When reproductive success is constant in one breeding phase, different tactics that increase variation in reproductive success among individuals may evolve in other phases. For instance, in shorebirds, which usually have a limited clutch size of four eggs, variation in reproductive tactics among individuals is expected either before egg-laying (e.g. diverse mating systems) or after hatching of the young (e.g. diverse parental care). In this paper, I studied the pied avocet (Recurvirostra avosetta), a shorebird with a modal clutch size of four eggs, to test whether post-hatch chick adoption as an alternative tactic can be linked to increased variation in annual reproductive success. When predation was high, naturally adopting pairs produced more filial fledglings than did pairs not adopting chicks and not losing chicks to adoption. The number of filial fledglings increased with the number of adopted young, possibly through diluting the chances of predation on filial young. Experimental chick addition did not lead to more fledged young due to low brood integrity as shown by the frequent loss of chicks from some experimental broods. When predation was low, larger broods occupied feeding territories with higher prey abundance than smaller broods, possibly due to their dominance over smaller ones. Pairs that lost chicks to adoption (donors) fledged as many filial young in their broods as did non-adopters/non-donors, whereas the total number of donors’ filial fledglings, including those raised in adopting broods, approached that of adopters. These findings show, for the first time, that post-hatch alternative reproductive tactics can lead to variation in annual reproductive success and to higher success for some pairs even in species where past adaptations limit variation in reproductive success in a certain phase of reproduction.

Estimation of Effective Population Size for the Long-Lived Darkblotched Rockfish Sebastes crameri

We report the variance effective population size (Ne) in darkblotched rockfish (Sebastes crameri) utilizing the temporal method for overlapping generations, which requires a combination of age-specific demography and genetic information from cohorts. Following calculations of age-specific survival and reproductive success from fishery data, we genotyped a sample (n = 1087) comprised by 6 cohorts (from 1995 to 2000) across 7 microsatellite loci. Our Ne estimate (Ne) plus 95% confidence interval was (Ne) = 9157 [6495-12 215], showing that the breeding population number could be 3-4 orders of magnitude smaller than the census population size (N) = 24 376 210). Our estimates resemble closely those found for fishes with similar life history, suggesting that the small (Ne)/(N) ratio for S. crameri is most likely explained by a combination of high variance in reproductive success among individuals, genetic structure, and demographic perturbations such as historical fishing. Because small (Ne)/(N) ratios have been commonly associated with potential loss of genetic variation, our estimates need careful consideration in rockfish management and conservation.

Pollen Limitation and Spatial Variation of Reproductive Success in the Insect-pollinated Shrub Chuquiraga Oppositifolia (Asteraceae) in the Chilean Andes

ABSTRACTLow temperatures, short growing seasons, and strong winds, which constrain the abundance and activity of insect pollinators, characterize alpine ecosystems. In northern hemisphere alpine environments, the reproductive output of several insect-pollinated plants has been reported to be pollen-limited. Using a supplemental hand-pollination experiment, we assessed the magnitude of pollen limitation (PL) in the obligate outcrossing insect-pollinated shrub Chuquiraga oppositifolia in the lower alpine scrub vegetation belt in the central Chilean Andes. We also assessed spatial variation in its reproductive success by comparing seed production among three additional sites. Hand-pollination resulted in a two- to three-fold increase in seed output above natural levels, thus demonstrating PL in this species. Nevertheless, percentage seed output remained low, increasing from 2.0 to 5.7%. Seed weight was reduced by 15% in hand-pollinated plants. Seed output was also low in the three additional sites, but did not differ among them, suggesting that low seed output is a spatially widespread phenomenon in this species. Our results, together with previous research, suggest that both pollinator visitation and abiotic resources (soil nutrients) constrain the reproductive output of this shrub. Longer-term research should unravel potential future vegetative and reproductive costs of current-year enhanced reproductive output and determine if PL is a recurrent phenomenon in the Chilean Andes.

Filial cannibalism as a conditional strategy in males of a paternal mouthbrooding fish

Filial cannibalism is widespread in a variety of animal species and has been generally accepted as an adaptive behavior. Within a population, some individuals adopt filial cannibalism and others do not, in spite of its adaptiveness. There is little knowledge of how such a polymorphic trait is maintained in nature. To understand the underlying mechanism of cannibalistic polymorphism, we conducted a long-term field study that involved monitoring of the reproductive experience of marked individuals in the paternal mouthbrooding cardinalfish, Apogon doederleini, in which parental males sometimes cannibalize their entire broods. We assumed that filial cannibalism can be described as one of three possible strategies: alternative, mixed or conditional. Individual cannibalistic tendencies, represented by the number of entire brood cannibalism performed by each individual in one breeding season, showed a random distribution within the study population. Moreover, the individual cannibalistic tendencies were not consistent between two successive seasons. These results suggest that filial cannibalism is phenotypically plastic, thus eliminating the alternative strategy as a possible mechanism. Comparison of variance in reproductive success between cannibals and non-cannibals showed that observations were not in accordance with those expected in the case that males adopt filial cannibalism stochastically, that is, as a mixed strategy. Our previous studies have indicated that filial cannibalism is affected by male status, such as age, somatic condition and mate availability. In conclusion, filial cannibalism by male A. doederleini is carried out as a conditional strategy.

Temporal recruitment patterns and gene flow in kelp rockfish (Sebastes atrovirens)

Pelagic dispersal of marine organisms provides abundant opportunity for gene flow and presumably inhibits population genetic divergence. However, ephemeral, fine-scale, temporal and spatial genetic heterogeneity is frequently observed in settled propagules of marine species that otherwise exhibit broad-scale genetic homogeneity. A large variance in reproductive success is one explanation for this phenomenon. Here, genetic analyses of 16 microsatellite loci are used to examine temporal patterns of variation in young-of-year kelp rockfish (Sebastes atrovirens) recruiting to nearshore habitat in Monterey Bay, California, USA. Population structure of adults from central California is also evaluated to determine if spatial structure exists and might potentially contribute to recruitment patterns. Genetic homogeneity was found among 414 young-of-year sampled throughout the entire 1998 recruitment season. No substantial adult population structure was found among seven populations spanning 800 km of coastline that includes the Point Conception marine biogeographic boundary. Comparison of young-of-year and adult samples revealed no genetic differentiation and no measurable reduction in genetic variation of offspring, indicating little variance in reproductive success and no reduction in effective population size for this year class. Simulation analyses determined that the data set was sufficiently powerful to detect both slight population structure among adults and a small reduction in effective number of breeders contributing to this year class. The findings of high gene flow and low genetic drift have important implications for fisheries management and conservation efforts.

Analysis of spatial genetic structure in an expanding Pinus halepensis population reveals development of fine‐scale genetic clustering over time

We analysed the change of spatial genetic structure (SGS) of reproductive individuals over time in an expanding Pinus halepensis population. To our knowledge, this is the first empirical study to analyse the temporal component of SGS by following the dynamics of successive cohorts of the same population over time, rather than analysing different age cohorts at a single time. SGS is influenced by various factors including restricted gene dispersal, microenvironmental selection, mating patterns and the spatial pattern of reproductive individuals. Several factors that affect SGS are expected to vary over time and as adult density increases. Using air photo analysis, tree-ring dating and molecular marker analysis we reconstructed the spread of reproductive individuals over 30 years beginning from five initial individuals. In the early stages, genotypes were distributed randomly in space. Over time and with increasing density, fine-scale (< 20 m) SGS developed and the magnitude of genetic clustering increased. The SGS was strongly affected by the initial spatial distribution and genetic variation of the founding individuals. The development of SGS may be explained by fine-scale environmental heterogeneity and possibly microenvironmental selection. Inbreeding and variation in reproductive success may have enhanced SGS magnitude over time.

Demographic and Genetic Estimates of Effective Population and Breeding Size in the AmphibianRana temporaria

Genetic methods for estimating effective population size ( Ne) or the effective number of breeders ( Nb) have become popular, but comparisons of these estimates with demographic estimates of Ne and Nb are rare, especially in anurans. We used three genetic (linkage disequilibrium, temporal moments, Bayesian coalescent-based method) and three demographic models, the latter considering number of breeding individuals, sex ratio, reproductive skew, and other demographic data, to estimate Ne and Nb in two subarctic populations (T and P) of the common frog Rana temporaria, subject to long-term capture-recapture studies. Demographic estimates of Ne based on total population size ( Ne ([T])= 44.5-56.9; Ne ([P])= 68.8-93.7) deviated markedly from the genetic estimates obtained using the linkage disequilibrium method ( Ne ([T])= 97.1; Ne ([P])= 13.2). The demographic estimates of Nb, taking into consideration sex ratio and variance in reproductive success ( Nb ([T])= 10.1-39.7; Nb ([P])= 3.9-21.3), were higher than the genetic estimates ( Nb ([T])= 3.7-5.4; Nb ([P])= 3.5-3.9). The main factors affecting the effective size estimates were sex ratio and reproductive skew. The discrepancies between corresponding Ne and Nb estimates highlight the sensitivity of both demographic and genetic estimates on their underlying assumptions. Yet the ratios of effective or breeding effective size to the census population size were similar to those reported earlier for anurans, reinforcing the view that the discrepancy between actual and effective breeding sizes in anuran populations is typically very large.

Effective size, census size, and genetic monitoring of the endangered razorback sucker, Xyrauchen texanus

For the past several decades the Lake Mohave population of the federally endangered razorback sucker, Xyrauchen texanus, has had no natural recruitment because of intensive predation on larvae by non-native fishes. In response to impending extirpation, a repatriation program was implemented where larval razorbacks are collected from the wild following natural spawning, reared in protective custody, and then repatriated at a much larger size. In this study, we estimated annual (Nbf) and generational female effective size (Nef) of the spawning stock by characterizing temporal genetic changes in mtDNA among larval cohorts, and then compared these estimates to the estimated number of reproductively capable (adult) females in Lake Mohave (Nf). Razorback suckers have life history and mortality schedules that could yield values of Nef/Nf as low as 10−5 due to match–mismatch recruitment (or the “Hedgecock effect”) that increases variance in reproductive success. Average Nbf was estimated to be 160, Nef was 706, and the ratio Nef/Nf was 0.29 and 0.38 for arithmetic and harmonic mean Nf, respectively. Our findings indicate that (i) larval sampling in Lake Mohave sufficiently encompasses temporal and spatial variation within annual larval cohorts as to be representative of the spawning stock, (ii) roughly 3–16% (about 8% on average) of adult females contribute genetically to larval cohorts each year, and (iii) repatriated fishes appear to be contributing genetically to larval cohorts. Simultaneous genetic and demographic monitoring offers insights that neither approach can provide alone into effects of population decline and management practices in this species.

The effects of isolation and colonization history on the genetic structure of marine-relict populations of Atlantic cod (Gadus morhua) in the Canadian Arctic

The genetic consequences of extended periods at low population size are fundamental to the conservation of depleted species such as the Atlantic cod (Gadus morhua). We compared microsatellite genetic variability among cod populations in Canadian Arctic lakes with that of Gilbert Bay resident and inshore cod from eastern Canada. The Arctic populations had the lowest genetic diversity and were the most strongly genetically structured and distinct. By contrast, eastern Canadian samples expressed high allelic diversity and were not significantly genetically structured or distinct relative to each other, whereas Gilbert Bay resident cod were intermediate to the Arctic and eastern Canadian groups. Our results are consistent with the hypothesis that the Arctic populations were colonized between 8000 and 5000 years ago and have experienced little or no gene flow since that time. Despite isolation at the extreme of the species' range, the Arctic populations have retained relatively high heterozygosities and high genetic effective population sizes relative to census sizes (Ne–Ncratios). Potential explanations for this include the absence of fishing pressure, allowing for the persistence of large, highly fecund individuals, as well as biotic (e.g., absence of planktivores) and abiotic (e.g., low environmental stochasticity) factors in the Arctic lakes that minimize individual variance in reproductive success.

Mating system and reproductive success in eastern Pacific harbour seals

Harbour seals sometimes breed along inland travel corridors where females become clustered in space and time and males establish underwater acoustic display territories similar to terrestrial arenas known as resource-based leks. Under these conditions, we predicted that higher levels of polygyny would be observed than has been previously reported for this species mating in open coast environments without travel corridors. Reproductive success (RS) of 70 males was measured using 20 microsatellite DNA loci and likelihood-based paternity analysis of 136 offspring collected over 3 years. Most males were assigned either zero or one paternity with 80% confidence. The greatest number of pups assigned to one male in a season was two. Variance in RS was higher for males than females (which are biologically limited to one offspring per year) indicating low to mild polygyny. In addition, distributions of relatedness values among pups within year classes did not differ significantly from a simulated distribution with R = 0, indicating that half-siblings were uncommon. Overall, polygyny levels were low relative to terrestrial pinniped mating systems and similar to observations from a harbour seal population along an open coast. Due to large confidence intervals associated with our results, we cannot rule out the hypothesis that a travel corridor might increase the degree of polygyny skew relative to that observed in open coast environments. Habitat appeared to influence male strategies as the most successful males in open coast environments patrolled offshore, while the most successful male in this study defended a territory along the travel corridor.

Comparative demography of treecreepers: evaluating hypotheses for the evolution and maintenance of cooperative breeding

Despite the long history of research on cooperative breeding, few comparative studies have been undertaken to test hypotheses for the evolution and maintenance of delayed dispersal using data from both cooperative and noncooperative species. We tested predictions about demographic differences between cooperative and noncooperative species based on four hypotheses for the evolution of delayed dispersal: the ecological constraints hypothesis, the life history hypothesis, the broad constraints hypothesis and the benefits-of-philopatry hypothesis. We controlled for phylogeny and habitat by comparing sympatric populations of two Australasian treecreepers (Climacteridae): the cooperatively breeding brown treecreeper, Climacteris picumnus, and the pair-breeding white-throated treecreeper, Cormobates leucophaea, which has lost cooperative breeding despite phylogenetic conservation of the trait in most of the treecreeper lineage. Our data failed to support nearly every prediction of the constraints and life history hypotheses. The two species had similar levels of constraints and similar life histories, and any significant differences were in the opposite direction to those predicted. However, the cooperative species was characterized by significantly higher variance in reproductive success as a result of group size effects, supporting the benefits-of-philopatry hypothesis as an explanation for the maintenance of delayed dispersal. Our comparisons also suggest that the benefits parents derive from group living determine whether or not they tolerate offspring on the natal territory. We therefore propose an antipredator tactics hypothesis in which delayed dispersal will initially evolve or be lost in particular taxa depending on the relative costs and benefits of vigilance and active defence versus concealment-based antipredator tactics.