Fitness consequences of male parental care in Savannah sparrows

The fitness consequences of male parental care are central tolife-history models of reproductive effort and the evolutionof social and genetic mating system However in a 4-year study(1992–1995) of Savannah sparrows (Parsserculs sandwichensis)nesting on Kent Island, New Brunswick, Canada, male parentalcare during the nestling stage was only weakly related to offspringquality and survivorship. Females compensated for poor paternalperformance by elevating their feeding rates, and they thusameliorated selection on male parental care. The likelihoodand size of subsequent broods was also unrelated to male feedingeffort, suggesting the absence of some indirect (or ‘female-mediated’)benefits. These results were confirmed by an experiment in whichmale removal resulted in increased female feeding rates buthad little effect on the number or quality of young. In contrast,previous research with multilocus DNA fingerpnnting revealeda strong, positive relationship between a male's parental effortand his subsequent fertilization success. Together, this researchsuggests that the proximate and ultimate targets of male feedingeffort can be disiinct traditional assumptions regarding thefunction of male parental care in birds may therefore be incomplete.     

Extra-pair paternity in monogamous and polygynous Savannah sparrows,Passerculus sandwichensis

Extra-pair paternity can influence mating systems by affecting the fitness costs associated with polygyny. Polygyny is disadvantageous to males when the time and energetic demands of multiple pairings decrease either a male's success at gaining extra-pair fertilizations or his ability to ensure paternity among harem members. In Savannah sparrows,Passerculus sandwichensison Kent Island, New Brunswick, Canada, multilocus DNA fingerprinting of 136 adults and young revealed substantial female infidelity: overall, 31 of 92 young (33.7%) in 15 of 24 nests (62.5%) were the product of extra-pair fertilizations. Male mating status was a strong predictor of paternity. Each of seven monogamous females produced at least one extra-pair offspring, but only six of 11 primary females (54.5%) and two of six secondary females (33.3%) were unfaithful. As a result, nearly 80% of the young in nests of polygynous males resulted from within-pair fertilizations, compared with only 40% of the young in nests of monogamous males. Kent Island Savannah sparrows are simultaneously polygynous, and the absence of paternity costs associated with polygyny is surprising. The observed pattern of paternity suggests the operation of female choice, although male control of parentage cannot be excluded.     

Climate and body size influence nest survival in a fish with parental care

1. The current study examined the effect of broad-scale climate and individual-specific covariates on nest survival in smallmouth bass over a 20-year period. 2. Large-scale climate indices [winter North Atlantic Oscillation (NAO) and winter El Ni?o/Southern Oscillation (ENSO)] and body size of parental males were important covariates in nest survival along with nest age and a quadratic trend in survival. 3. We did not find an effect due to a habitat covariate (total effective fetch) or a phenology covariate (degree-days at start of nesting) on nest survival. 4. Male size in the second half of the nesting season was a more influential covariate on nest success than male size in the first half or throughout the nesting period. 5. We present evidence showing that winter NAO/ENSO indices establish limnological conditions the following spring that influence thermal stability of the lake during the nesting period. 6. The combined climate and body size covariates point to nest survival as a function of lagged climate-scale influences on limnology and the individual-scale influence of bioenergetics on the duration of parental care and nest success.     

Stress,song and survival in sparrows

The stress response—increases in circulating glucocorticoids following a stressor—is typically considered adaptive, but few studies address the fitness consequences of individual variation in stress response. Generally, due to negative consequences of prolonged elevation of glucocorticoids, animals should have a transient stress response just sufficient to cope with the stressor. In rodents, stress responsiveness is affected by early developmental experience, and hyper-responsiveness to stress is linked to morbidity and mortality. We assessed individual variation in stress responses in free-living song sparrows, Melospiza melodia, in relation to fitness-related measures including song and overwinter survival. Birds with greater increases in corticosterone 30 min following restraint stress were less likely to return to breed the following year. Stress responsiveness was also correlated with song complexity: males with fewer syllables in their song repertoires had greater stress reactivity. Our findings support the hypothesis that developmental stressors both impair song development and affect the adult stress response. Thus, individual variation in the stress response may relate to variation in fitness.     

Postfledging parental care and unexpected foraging behaviour of juvenile Sooty Terns Onychoprion fuscata in the southern Mozambique Channel

The Berlese funnel method, and its variations, is commonly used for the extraction of arthropods from various substrates such as nest material. However, little is known about its effectivity in extracting ectoparasites from penguin nests. This study aimed to assess the efficacy of a modified Berlese funnel method in extracting ectoparasites from African Penguin Spheniscus demersus nests and to determine differences in extraction capacity across parasite taxa and life stages. African Penguin nests (n = 278) were sampled at five penguin colonies along the south-west coast of South Africa in 2017. Nest samples were subjected to a modified Berlese funnel method with naphthalene as a repellent. Thereafter, all remaining ectoparasites were removed by hand sorting. Ectoparasite abundance was positively correlated between the two extraction methods (Spearman’s r = 0.73–0.89). Compared to total counts, flea (combined life stages and larvae) and total ectoparasite abundance and prevalence were significantly lower with the modified Berlese funnel method. In addition, adult flea and tick prevalence (but not abundance) was significantly lower with the modified Berlese funnel method compared with total counts. The modified Berlese funnel method fails as a quantitative method and can only provide a crude indication of the incidence of ectoparasites in penguin nests.     

Propagule size and parental care: the safe harbor hypothesis

Species that protect their offspring (by egg-brooding or live-bearing) produce larger (and hence fewer) propagules than do related species that do not protect their offspring. This result holds true in most animal groups showing parental care. Analysis of these data supports the hypothesis that propagule size evolves in response to relative survivorship rates during and after the propagule stage: natural selection is most likely to favor large egg size when the egg is a safe harbor.     

Genetic similarity, extrapair paternity, and offspring quality in Savannah sparrows (Passerculus sandwichensis)

The occurrence of extrapair paternity (EPP) in birds is oftenattributed to the action of good-genes sexual selection wherebyfemales trade up on male genetic quality by allocating fertilizationsto males with better genes than those possessed by their socialmate. To date, most studies of EPP in birds focus on absolutemeasures of male quality as a criterion for female choice, althoughmultiple mating by females in other taxa is more commonly ascribedto benefits associated with the individual optimization of offspringgenotypes. Here, we examine whether the genetic similarity ofsocial mates predicts female mating patterns in a populationof Savannah sparrows (Passerculus sandwichensis) where as manyas 70% of adults produce extrapair young (EPY). We considerthe influence of genetic similarity across all stages of a female'sdecision-making process, from pair formation through the decisionto produce EPY, to the allocation of fertilizations to specificextrapair sires. In a 4-year study of 175 males, 206 females,and 506 offspring, females were more likely to produce EPY whenpaired to genetically similar males, but they did not appearto be influenced by the size, age, mass, individual heterozygosity,and genetic diversity of their social mates. In paired comparisons,females were almost twice as likely to decrease their geneticsimilarity to males when producing EPY as they were to increaseit. Nonetheless, females did not select especially dissimilarmales when mating outside the pair-bond nor did they pair disassortativelywith respect to genetic similarity. Relative measures of malequality may influence mating patterns in birds, but only atsome points in a female's decision-making process.     

Determinants of parental care and offspring survival during the post-fledging period: males care more in a species with partially reversed sex roles

Sexual conflict is magnified during the post-fledging period of birds when the sexes face different trade-offs between continuing parental care or investing in self maintenance or other mating opportunities. Species with reversed sex roles provide a unique opportunity to study the relationship between mating systems and investment in parental care. Here, we provide the first detailed study of the length of care by males versus females (n = 24 pairs) during the post-fledging period, assessing factors that may promote care within and between the sexes. In the northern flicker Colaptes auratus, a species with partly reversed sex roles, males cared longer than females (average 16 versus 12 days, respectively). Overall, 36 % of females but no males deserted the brood prior to fledgling independence. Parents that provisioned nestlings at a high rate also spent more days feeding fledglings. Among males, age and nestling feeding rates were positively associated with the length of care. Among females, a low level of feather corticosterone (CORT_f) was associated with a longer length of care. About 45 % of fledglings died within the first week, but fledglings with intermediate body mass had the highest survival suggesting stabilizing selection on mass. Fledgling survival was also higher in individuals with larger broods and lower levels of CORT_f. We demonstrate that because females can be polyandrous they often desert the brood before males, and that the sexes respond to different cues relating to their energy balance when deciding the length of care given to their offspring.     

Sexual dimorphism,survival, and parental investment in relation to offspring sex in a precocial bird

Differential growth rate between males and females, owing to a sexual size dimorphism, has been proposed as a mechanism driving sex‐biased survival. How parents respond to this selection pressure through sex ratio manipulation and sex‐biased parental investment can have a dramatic influence on fitness. We determined how differential growth rates during early life resulting from sexual size dimorphism affected survival of young and how parents may respond in a precocial bird, the black brant Branta bernicla nigricans. We hypothesized that more rapidly growing male goslings would suffer greater mortality than females during brood rearing and that parents would respond to this by manipulating their primary sex ratio and parental investment. Male brant goslings suffered a 19.5% reduction in survival relative to female goslings and, based on simulation, we determined that a female biased population sex ratio at fledging was never overcome even though previous work demonstrated a slight male‐biased post‐fledging survival rate. Contrary to the Fisherian sex ratio adjustment hypothesis we found that individual adult female brant did not manipulate their primary sex ratio (50.39% male, n = 645), in response to the sex‐biased population level sex ratio. However, female condition at the start of the parental care period was a good predictor of their primary sex ratio. Finally, we examined how females changed their behavior in response to primary sex ratio of their broods. We hypothesized that parents would take male biased broods to areas with increased growth rates. Parents with male biased primary sex ratios took broods to areas with higher growth rates. These factors together suggest that sex‐biased growth rates during early life can dramatically affect population dynamics through sex‐biased survival and recruitment which in turn affects decisions parents make about sex allocation and sex‐biased parental investment in offspring to maximize fitness.     

The evolutionary puzzle of egg size,oxygenation and parental care in aquatic environments

Offspring fitness generally improves with increasing egg size. Yet, eggs of most aquatic organisms are small. A common but largely untested assumption is that larger embryos require more oxygen than they can acquire through diffusion via the egg surface, constraining egg size evolution. However, we found no detrimental effects of large egg size on embryo growth and survival under hypoxic conditions. We tested this in the broad-nosed pipefish, Syngnathus typhle, whose males provide extensive care (nourishment, osmoregulation and oxygenation) to their young in a brood pouch on their bodies. We took advantage of this species'' pronounced variation in egg size, correlating positively with female size, and tested the effect of hypoxia (40% dissolved oxygen) versus fully oxygenated (100%) water on embryo size and survival of large versus small eggs after 18 days of paternal brooding. Egg size did not affect embryo survival, regardless of O₂ treatment. While hypoxia affected embryo size negatively, both large and small eggs showed similar reductions in growth. Males in hypoxia ventilated more and males with large eggs swam more, but neither treatment affected their position in the water column. Overall, our results call into question the most common explanation for constrained egg size evolution in aquatic environments.     

Effects of parental survival on clutch size decisions in fluctuating environments

Whereas in constant environments parental survival has no effect on optimal clutch size in the absence of trade-offs between juvenile and parental survival, the situation is drastically different in fluctuating environments. We consider a model in which, with respect to reproduction, parents and offspring are equivalent at the start of the next breeding season. When generations are non-overlapping, the clutch size maximizing geometric mean surviving number of offspring is optimal among all pure clutch size strategies. We prove that, as parental survival increases relative to that of the offspring, the optimal clutch size converges to the arithmetic mean maximizing clutch size (the so-called ‘Lack clutch size’). We also give a numerical procedure for calculating optimal mixed strategies and we show that, as environmental variance increases and/or parental survival decreases, mixed rather than pure strategies become optimal. Furthermore, we explain how to estimate fitness from empirical data under the assumptions of our model. This revised version was published online in July 2006 with corrections to the Cover Date.     

The evolution of parental care and egg size: a comparative analysis in frogs

The evolution of parental care and egg size has attracted considerable attention and theoretical debate. Several different hypotheses have been proposed concerning the trajectories of parental care and egg size evolution and the order of specific evolutionary transitions. Few comparative studies have investigated the predictions of these hypotheses. Here, we investigate the evolutionary association between parental care and egg size in frogs in a phylogenetic context. Data on egg size and presence or absence of parental care in various species of frogs was gathered from the scientific literature. As a basis for our comparative analyses, we developed a phylogenetic supertree, by combining the results of multiple phylogenetic analyses in the literature using matrix representation parsimony. Using phylogenetic pairwise comparisons we demonstrated a significant association between the evolution of parental care and large egg size. We then used recently developed maximum likelihood methods to infer the evolutionary order of specific transitions. This analysis revealed that the evolution of large egg size typically precedes the evolution of parental care, rather than the reverse. We discuss the relevance of our results to previous hypotheses concerning the evolution of parental care and egg size.     

Birth order, individual sex and sex of competitors determine the outcome of conflict among siblings over parental care

Success in competition for limiting parental resources depends on the interplay between parental decisions over allocation of care and offspring traits. Birth order, individual sex and sex of competing siblings are major candidates as determinants of success in sib-sib competition, but experimental studies focusing on the combined effect of these factors on parent-offspring communication and within-brood competitive dynamics are rare. Here, we assessed individual food intake and body mass gain during feeding trials in barn swallow chicks differing for seniority and sex, and compared the intensity of their acoustic and postural solicitation (begging) displays. Begging intensity and success in competition depended on seniority in combination with individual sex and sex of the opponent. Junior chicks begged more than seniors, independently of satiation level (which was also experimentally manipulated), and obtained greater access to food. Females were generally weaker competitors than males. Individual sex and sex of the opponent also affected duration of begging bouts. Present results thus show that competition with siblings can make the rearing environment variably harsh for developing chicks, depending on individual sex, sex of competing broodmates and age ranking within the nest. They also suggest that parental decisions on the allocation of care and response of kin to signalling siblings may further contribute to the outcome of sibling competition.     

Sexual selection and the geography of Plasmodium infection in Savannah sparrows (Passerculus sandwichensis)

According to Hamilton and Zuk's hypothesis of parasite-mediated sexual selection, host-parasite coevolution maintains variation in male genetic quality and allows for strong intersexual selection in species with high rates of infection. In birds, most interspecific tests of this hypothesis relate the prevalence of blood parasites to some measure of the intensity of sexual selection. Such tests often rely on limited sampling of single populations to estimate species-wide infection rates, and many tests are thus vulnerable to intraspecific (geographic) variation in the evolutionary ecology of disease. Here, we used molecular techniques to examine variation in the prevalence of Plasmodium spp. across 14 populations of Savannah sparrows, Passerculus sandwichensis, in eastern North America. Plasmodium could not be detected in any of 68 island birds, but 34 of 119 (29%) mainland males, and 7 of 43 (16%) mainland females were infected. Among mainland birds, infection was common in southern populations but rare in New Brunswick, Canada. Overall, the prevalence of Plasmodium ranged from 0 to 60% across populations, although only 17.8% of birds were infected in the pooled (species-wide) sample. The extent of this geographic variation suggests that limited sampling of single populations is unlikely to yield accurate estimates of species-wide infection rates. However, among mainland Savannah sparrows, the prevalence of malaria correlated strongly with average male size and the degree of sexual size dimorphism. We speculate that either sexual selection leads to male-biased infection or, conversely, that high rates of infection promote the evolution of strong intersexual selection.     

Offspring fitness varies with parental extra-pair status in song sparrows, Melospiza melodia

Numerous studies have tested for indirect selection on female extra-pair reproduction (EPR) by quantifying whether extra-pair young (EPY) are fitter than their within-pair young (WPY) maternal half-siblings. In contrast, the hypothesis that offspring of EPY and WPY (rather than the EPY and WPY themselves) differ in fitness has not been tested, even though inter-generational effects of parental extra-pair status on offspring fitness could alter the magnitude and direction of indirect selection on EPR. We tested whether offspring of EPY song sparrows, Melospiza melodia, were more likely to recruit or produce hatched or recruited offspring over their lifetimes than offspring of WPY. Hatchlings with one or two EPY parents were more likely to recruit and produce hatched offspring than hatchlings with two WPY parents. Furthermore, these relationships differed between maternal versus paternal extra-pair status. Hatchlings with EPY fathers were more likely to recruit and produce offspring than hatchlings with WPY fathers. In contrast, hatchlings with EPY mothers were as likely to recruit as hatchlings with WPY mothers and tended to be less likely to produce recruited offspring. Depending on the causal genetic and environmental mechanisms, such conflicting inter-generational relationships between parental extra-pair status and offspring fitness could substantially influence the evolutionary dynamics of EPR.     

Evidence for calibration of magnetic migratory orientation in Savannah sparrows reared in the field

The orientation system of migratory birds consists of a magnetic compass and compasses based upon celestial cues. In many places, magnetic compass directions and true or geographic compass directions differ (referred to as magnetic declination). It has been demonstrated experimentally in several species that the innate preferred direction of magnetic orientation can be calibrated by celestial rotation, an indicator of geographic directions. This calibration process brings the two types of compass into conformity and provides the birds with a mechanism that compensates for the spatial variation in magnetic declination. Calibration of magnetic orientation has heretofore been demonstrated only with hand-raised birds exposed to very large declination (90° or more). Here we show that the magnetic orientation of wild birds from near Albany, New York, USA (declination = 14° W) was N–S, a clockwise shift of 26° from the NNW–SSE direction of birds raised entirely indoors. Hand-raised birds having visual experience with either the daytime sky or both day and night sky orientated N–S, similar to wild-caught birds. These data provide the first confirmation that calibration of magnetic orientation occurs under natural conditions and in response to modest declination values.     

Sexual conflict over parental care promotes the evolution of sex differences in care and the ability to care

Strong asymmetries in parental care, with one sex providing more care than the other, are widespread across the animal kingdom. At present, two factors are thought to ultimately cause sex differences in care: certainty of parentage and sexual selection. By contrast, we here show that the coevolution of care and the ability to care can result in strong asymmetries in both the ability to care and the level of care, even in the absence of these factors. While the coevolution of care and the ability to care does not predict which sex evolves to care more than the other, once other factors give rise to even the slightest differences in the cost and benefits of care between the sexes (e.g. differences in certainty in parentage), a clear directionality emerges; the sex with the lower cost or higher benefit of care evolves both to be more able to care and to provide much higher levels of care than the other sex. Our findings suggest that the coevolution of levels of care and the ability to care may be a key factor underlying the evolution of sex differences in care.