Quantifying the costs and benefits of parental care in female treehoppers

Parental protection of eggs represents one of the most basicforms of parental care. Theory suggests that even such basicparental investment represents a trade-off between current offspringsurvival and future reproductive success. However, few studieshave quantified the underlying costs and benefits of parentalcare for marked individuals across an entire lifetime. I markedand followed 370 females of Publilia concava (Hemiptera: Membracidae)that exhibited a range of guarding durations for their firstclutch. Greater hatching success was correlated with longerguarding durations, and a removal experiment verified that femalepresence was responsible for a twofold increase in hatchingsuccess. On the other hand, females that remained to guard eggshad a lower number and size of future broods, suggesting thatparental care may reduce lifetime fecundity. Marked femalesexhibited a bimodal distribution of guarding durations, reflectingthe extreme tactics of immediate abandonment or remaining throughhatching. Estimates of lifetime number of nymphs produced byfemales that abandon eggs early versus guard eggs through hatchingrevealed roughly equivalent levels of fitness. I discuss theconditions under which we might expect a female to adopt eachof the alternative tactics, given the costs and benefits ofparental care that were quantified in this study.     

Risk taking during parental care: a test of the harm-to-offspring hypothesis

Amount of risk taking during parental care is often explainedin relation to the reproductive value of the offspring. The"harm-to-offspring hypothesis" focuses on the relative harma period of no parental care can do to the offspring. Accordingto this hypothesis, parents should take greater risks for offspringin poor condition than for offspring in good condition. We manipulatedoffspring condition in the pied flycatcher (Ficedula hypoleuca)and tested the harm-to-offspring hypothesis by exposing parentsto a predator model (a sparrowhawk, Accipiter nisus). Time elapseduntil a parent first entered the nest-box was used as a risk-takingmeasure. Parents spent significantly shorter time until first nestvisit for offspring in poor condition than for offspring ingood condition. Hence, the harm-to-offspring hypothesis wassupported.     

Brood desertion by female shorebirds: a test of the differential parental capacity hypothesis on Kentish plovers.

The aim of this study was to examine whether the energetic costs of reproduction explain offspring desertion by female shorebirds, as is suggested by the differential parental capacity hypothesis. A prediction of the hypothesis is that, in species with biparental incubation in which females desert from brood care after hatching, the body condition of females should decline after laying to a point at which their body reserves are too low for continuing parental care. We tested this prediction on Kentish plovers (Charadrius alexandrinus) in which both sexes incubate but the females desert from brood care before the chicks fledge. We found no changes in either the body masses or body compositions of both individual male and female plovers from early incubation and throughout early chick rearing. Furthermore, the timing of brood desertion by females was not affected by their body condition. Neither did we find gender differences in the energetic costs of incubation. There were no differences in the timing of brood desertion between experimental and control females in an experiment in which we lengthened or shortened the duration of incubation by one week. These results indicate that energetic costs do not explain offspring desertion by female Kentish plovers and that the needs of chicks for parental care rather than cumulative investment by females is what determines the timing of brood desertion.     

Seasonal Variations in the Diet and Foraging Behaviour of Dunlins Calidris alpina in a South European Estuary: Improved Feeding Conditions for Northward Migrants

During the annual cycle, migratory waders may face strikingly different feeding conditions as they move between breeding areas and wintering grounds. Thus, it is of crucial importance that they rapidly adjust their behaviour and diet to benefit from peaks of prey abundance, in particular during migration, when they need to accumulate energy at a fast pace. In this study, we compared foraging behaviour and diet of wintering and northward migrating dunlins in the Tagus estuary, Portugal, by video-recording foraging birds and analysing their droppings. We also estimated energy intake rates and analysed variations in prey availability, including those that were active at the sediment surface. Wintering and northward migrating dunlins showed clearly different foraging behaviour and diet. In winter, birds predominantly adopted a tactile foraging technique (probing), mainly used to search for small buried bivalves, with some visual surface pecking to collect gastropods and crop bivalve siphons. Contrastingly, in spring dunlins generally used a visual foraging strategy, mostly to consume worms, but also bivalve siphons and shrimps. From winter to spring, we found a marked increase both in the biomass of invertebrate prey in the sediment and in the surface activity of worms and siphons. The combination of these two factors, together with the availability of shrimps in spring, most likely explains the changes in the diet and foraging behaviour of dunlins. Northward migrating birds took advantage from the improved feeding conditions in spring, achieving 65% higher energy intake rates as compared with wintering birds. Building on these results and on known daily activity budgets for this species, our results suggest that Tagus estuary provides high-quality feeding conditions for birds during their stopovers, enabling high fattening rates. These findings show that this large wetland plays a key role as a stopover site for migratory waders within the East Atlantic Flyway.     

Brood defense and brood size in the great tit (Parus major): a test of a model of unshared parental investment

To understand the interaction of the many contextual variablesthat affect parental behavior a number of static optimalitymodels have been developed. Among these the one by Lazarus andInglis (1986) is the only one to specifically predict the magnitudeof unshared parental investment (PI), i.e., of parental carethat carries a cost to the parent and that benefits all currentoffspring equally because it cannot be divided among them. Weinvestigated specifically how parent great tits (Parus major)gear their brood defense, a form of unshared PI, to the sizeof the brood at stake and to the risk incurred as a functionof the type of predator. The predators used were dummies ofthe great spotted woodpecker (Picoides major) and of the tawnyowl (Strix aluco). Normally, adults can approach the woodpeckerwith impunity; it had inflicted heavy losses to nestlings ofthe study populations of great tits near Wolfsburg, Lower Saxony.Parent great tits whose brood had been artificially reducedto two young responded to the dummy with less defense than dida control group with their pre-test brood size left intact.The nature of defense was qualitatively the same as that elicitedby a live woodpecker. Parents confronting the owl near theirbrood decreased their response with an artificial reductionin brood size much less. Because the owl used poses a seriousrisk to the defenders, as compared to the woodpecker, the resultlends powerful support to the "total loss" version of the modelof unshared PI; it predicts brood size to affect unshared PImore strongly when there is less risk to the parent. This interpretationis correct to the extent that one premise of the model, namelythat of uncompromised parentage, can be relaxed; great tit broodscontain a sizeable number of extrapair young. Males defendedtheir brood more strongly than did females. Sex and brood manipulationadded up linearly when affecting defense level, i.e., therewas no interaction.     

Spring temperature, clutch initiation date and duck nest success: a test of the mismatch hypothesis

1. Increases in average global temperature during the twentieth century have prompted calls for research on the effect of temperature variation on avian population dynamics. Particular attention has been paid to the hypothesis that increased temperatures may affect a species' ability to shift their breeding efforts to match the phenology of their prey, and thus result in reduced reproductive success (the 'mismatch hypothesis'). 2. We used data from a long-term study of breeding ducks to investigate how duck nest success varied with clutch initiation date, and to test whether spring temperature affected this relationship in a manner consistent with the mismatch hypothesis. We modelled five possible functional forms of how nest success might vary with clutch initiation date and spring temperature, and used an information-theoretic approach to determine which model best described the nesting outcomes of five dabbling duck species nesting in Saskatchewan, Canada. 3. Probability of nest success for the five species did not vary strongly with clutch initiation date, and we found evidence consistent with the mismatch hypothesis for one species, northern pintail Anas acuta, although weight of evidence was weak. 4. Overall nest success of all five species was positively associated with spring temperature. These results suggest that increasing spring temperature alone (within the range observed in this study) may not affect nest success in a manner that would result in lower populations of breeding ducks.     

Body size,duration of parental care,and the intrinsic rate of natural increase in eutherian and metatherian mammals

Summary The intrinsic rate of natural increase (r _m), conception to weaning time (t _cw), age of first reproduction (t_mat), and components of fecundity were compared between ecologically similar groups of 42 metatherian (=marsupial) and 42 eutherian mammals. Marsupial t _cw s average 50% longer than those of eutherians. Small marsupials (<400 g) mature later, and have lower and r _ms than eutherians; large marsupials (>10,000 g) do not mature later but also have lower r _ms. At body sizes of 1,000–3,500 g, marsupials and eutherians have similar t _mat s and t _cw s but marsupials have greater r _ms. Marsupials compensate for their longer t _cw s by a variety of methods including embryonic diapause, larger litter sizes, and short periods between weaning and maturity. Although the greatest similarities in marsupial and eutherian life histories are at body sizes of 1–5 kg, compensation for long t _cw may be seen at any marsupial body size. Other ecological factors not withstanding, marsupial reproduction is neither inherently inferior to that of eutherians nor obviously more advantageous in unpredictable environs.     

Predictable males and unpredictable females: sex difference in repeatability of parental care in a wild bird population

Repeatability of parental care, let alone heritability of care, has been rarely measured, although there has been much research linking sexual selection to male parental care and also examining biparental care in relation to game theory models. We investigated within- and between-year repeatabilities of incubation and nestling provisioning and how these two types of parental care were related in a sexually dimorphic species, the house sparrow, Passer domesticus. We found that between- and within-year repeatabilities of feeding rate were high in males and low to moderate in females, but that between- and within-year repeatabilities of incubation time were low to moderate in both sexes. Interestingly, the amount of time during which neither sex incubated significantly predicted the subsequent male feeding rate but not the female feeding rate. Our results suggest a need for a new theoretical framework that encompasses variation in the predictability and plasticity of parental investment by individuals.     

Body size and fecundity in the waterstrider Aquarius remigis: a test of Darwin's fecundity advantage hypothesis

The general female bias in body size of animals is usually attributed to fecundity selection. While many studies have demonstrated a positive relationship between body size and fecundity, the most common interpretation of fecundity selection is that larger females have larger abdomens and can hold more eggs, yet the relationship between abdomen size and fecundity has rarely been examined. For the waterstrider, Aquarius remigis, we find a significant relationship between body size and fecundity and demonstrate that the target of fecundity selection is abdomen size. Thus, larger females have higher fecundities because they have larger abdomens and not because of their total size per se. The rate at which fecundity increases with increasing abdomen size exceeds that which would be expected due to a simple volume constraint and suggests that other factors, such as increased ability to obtain resources, may contribute to the increase in fecundity with body size. Selection intensities estimated from our data indicate that fecundity selection could be a significant selective force on both total and abdomen lengths. Previous studies have found that abdomen size increased faster than body size and thus, larger females had relatively larger abdomens. The relationship of abdomen length and thorax length in A. remigis is hypoallometric and indicates that larger females have relatively smaller abdomens. We hypothesize that this may reflect conservation of abdomen size in females developing under poor conditions. Finally, while egg size is not directly related to body size, we find a trade-off between egg size and number when female abdomen length is held constant, suggesting that selection on egg size may influence abdomen length only indirectly through its effects on fecundity.     

Aspects of embryonic and postembryonic development of the Japanese freshwater crayfish Cambaroides japonicus (Crustacea, Decapoda) including a hypothesis on the evolution of maternal care in the Astacida

Little is known about the biology of the East Asian freshwater crayfish of the genus Cambaroides. Furthermore, the phylogenetic relationships of Cambaroides are controversial. To gain more information about East Asian crayfish and to shed some light on the phylogeny and evolution of freshwater crayfish, some aspects of the embryonic and postembryonic development of the Japanese freshwater crayfish species Cambaroides japonicus are described. The general appearance of the embryo and the growth zone consisting of about 40 ectoteloblasts correspond with the apomorphic pattern described for all other freshwater crayfish species. The occurrence of eight mesoteloblasts is a typical malacostracan character. In addition to the occurrence of freshwater crayfish apomorphies, such as a telson thread, the postembryonic development of C. japonicus follows the ground pattern of the Northern Hemisphere Astacoidea. In particular, some characteristics are in common with the postembryonic development of the Astacidae. These concern the shape of the telson of the juvenile stage 1 and the occurrence of plumose telson setae in juvenile stage 2. Furthermore, the recurved hooks of the hatchlings are lost in the juvenile stage 2, indicating an early independence of the juveniles from the mother. On the other hand, the early appearance of a spermatheca (annulus ventralis) is shared between C. japonicus and the other (American) Cambaridae. The hypothesis is developed that the evolutionary success of the different freshwater crayfish taxa might be related to the stepwise extension of maternal care as an adaptation to the freshwater environment.     

Faithful females receive more help: the extent of male parental care during incubation in relation to extra‐pair paternity in songbirds

Parental care provided by males occurs in a diverse array of animals and there are large differences among species in its extent compared with female care. However, social and ecological factors responsible for interspecific differences in male's share of parental duties remain unclear. Genetic fidelity of females has been long considered important. Theory predicts that females should receive more help from their mates in raising the offspring in species with high genetic fidelity. Using avian incubation behaviour as a model system, we confirmed this prediction. The extent of male's help during incubation increased with decreasing rate of extra‐pair paternity across species (22 species of socially monogamous songbirds from 13 families; male's share of incubation ranged from 6% to 58%), even after accounting for covariates, biases in species selection and intraspecific variability. Moreover, this result was not sensitive to two different phylogenies and branch length estimates. We suggest that our findings support the notion, backed by theory, that genetic fidelity is an important factor in the evolution of male parental care. We offer several behavioural scenarios for the coevolution between male's share of parental duties and the genetic mating system.     

Seasonal decline in provisioning effort and nestling mass of Blue Tits Cyanistes caeruleus: experimental support for the parent quality hypothesis

A common feature of many birds breeding in seasonal environments is that fitness‐related parameters such as nestling mass or survival decline as the breeding season progresses. Consequently, there is a tendency for early breeders to have better reproductive performance than individuals breeding later in the season. This variation could be caused by factors associated with the date of laying, such as changing environmental conditions (date hypothesis), or by differences in parental quality between early and late breeders (parent quality hypothesis). To evaluate the relative importance of both hypotheses, we manipulated hatch dates of Blue Tits Cyanistes caeruleus by exchanging clutches with different incubation stages and assessed the impact on nestling mass, nestling diet and provisioning rates. Mean nestling mass declined significantly over the season. This was the combined result of differences in parental quality, which dominated in the early part of the season, and the influence of hatching date (date effect per se), which prevailed later in the season. Nestling diet composition was apparently uninfluenced by the manipulation, suggesting that deteriorating food supplies are the primary reason for the seasonal variation in the nestling diet. Counter to the date hypothesis, delayed parents did not feed their young less than control pairs did, but in fact exhibited higher provisioning rates. Our results suggest that in this population, parental quality seems to constrain post‐hatching reproductive performance and such intrinsic limitations may help to explain why certain individuals breed later.     

The costs of parental care: a meta-analysis of the trade-off between parental effort and survival in birds

A fundamental premise of life-history theory is that organisms that increase current reproductive investment suffer increased mortality. Possibly the most studied life-history phenotypic relationship is the trade-off between parental effort and survival. However, evidence supporting this trade-off is equivocal. Here, we conducted a meta-analysis to test the generality of this tenet. Using experimental studies that manipulated parental effort in birds, we show that (i) the effect of parental effort on survival was similar across species regardless of phylogeny; (ii) individuals that experienced reduced parental effort had similar survival probabilities than control individuals, regardless of sex; and (iii) males that experienced increased parental effort were less likely to survive than control males, whereas females that experienced increased effort were just as likely to survive as control females. Our results suggest that the trade-off between parental effort and survival is more complex than previously assumed. Finally, our study provides recommendations of unexplored avenues of future research into life-history trade-offs.     

Feathers,suspicions, and infidelities: an experimental study on parental care and certainty of paternity in the blue tit

Theoretical models on parental care predict that males should decrease their parental effort when paternity is in doubt. Males may use some cues to assess their certainty of paternity, and try to avoid rearing offspring sired by extra‐pair males. We have previously reported in a socially monogamous passerine, the blue tit (Cyanistes caeruleus), that males decorate their nests with feathers, and that when this ornament is manipulated, males appear to have suspicions about the presence of an intruder male. Here, we decrease the male's certainty of paternity through experimental feather supplementation to analyse whether the outcome of our experiment supports the assumptions of the parental care theory. Male C. caeruleus responded to the feather supplementation experiment by reducing their parental investment (feeding frequency and nest defence) in comparison with control males. The occurrence of extra‐pair offspring in experimental nests was double than that in controls. This suggests that the manipulation was successful not only in altering males' perceived paternity, but also, indirectly, the actual paternity. Furthermore, males that gained extra‐pair young also had a higher than average probability to lose paternity in their nest, which may imply that male C. caeruleus faced a trade‐off between obtaining extra‐pair fertilizations and maintaining paternity in their own nest. Overall, this study supports the idea that males are prone to decrease their parental effort when they perceive that the risk of losing paternity is high. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 552–561.     

Culture and fertility in the Nepal Himalayas: A test of a hypothesis

This article tests the hypothesis that in the Nepal Himalayas the Tibetan sociocultural system (fraternal polyandry) reduces aggregate fertility whereas the Hindu system maximizes by early and virtually universal monogamous marriage. It is shown that while fraternal polyandry does reduce aggregate fertility, the hypothesis that it would lead to a substantially lower fertility rate is false because the effects of fraternal polyandry are roughly balanced by the effects of postwidowhood celibacy among the Hindus.This paper is the result of research undertaken in Nepal from 1975–1977. Support was provided in part by a National Science Foundation Dissertation Improvement Grant (BNS 76-00670) and grants from the Graduate Alumni Fund, Case Western Reserve University.     

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.     

Extrapair paternity in the great tit (Parus major): a test of the "good genes" hypothesis

In 1993 and 1994 we determined the frequency of extrapair paternityin broods of great tits, Parus major using multilocus DNA fingerprinting.We found no instances of intraspecific brood parasitism, but40% of broods (31/78) contained extrapair-fathered young and83% of offspring (58/681) were xtrapair We identified the geneticfathers of 60% of the extrapair nestlings (35/ 58). Males withfull and lost paternity did not differ significantly in traitsthat have been suggested to indicate male quality, nor did thegenetic and social fathers of extrapair offspring. In 1993,cuckolded males sired more offspring that recruited to the subsequentbreeding season than males with full paternity. Moreover, eventhough genetic fathers of extrapair young (EPY) sired more fledglingsthan the males they cuckolded, genetic and social fathers ofEPY did not differ in the number of recruits sired. Also, theEPY of a brood did not survive better than their half sibs.Thus, our results do not supportthe hypothesis that femaleschoose better quality males for extrapair matings ("good genes"hypothesis). Further, the level of extrapair paternity differedmarkedly between the two years. Our data show that females areconstrained in their extrapair activities by the availabilityof extrapair mates. This is at least partly due to yearly differencesin breeding synchrony.