Differential investment and costs during avian incubation determined by individual quality: an experimental study of the common eider (Somateria mollissima)

Individuals of different quality may have different investment strategies, shaping responses to experimental manipulations, thereby rendering the detection of such patterns difficult. However, previous clutch-size manipulation studies have infrequently incorporated individual differences in quality. To examine costs of incubation and reproductive investment in relation to changes in clutch size, we enlarged and reduced natural clutch sizes of four and five eggs by one egg early in the incubation period in female common eiders (Somateria mollissima), a sea duck with an anorectic incubation period. Females that had produced four eggs (lower quality) responded to clutch reductions by deserting the nest more frequently but did not increase incubation effort in response to clutch enlargement, at the cost of reduced hatch success of eggs. Among birds with an original clutch size of five (higher quality), reducing and enlarging clutch size reduced and increased relative body mass loss respectively without affecting hatch success. In common eiders many females abandon their own ducklings to the care of other females. Enlarging five-egg clutches led to increased brood care rate despite the higher effort spent incubating these clutches, indicating that the higher fitness value of a large brood is increasing adult brood investment. This study shows that the ability to respond to clutch-size manipulations depends on original clutch size, reflecting differences in female quality. Females of low quality were reluctant to increase investment at the cost of lower hatch success, whereas females of higher quality apparently have a larger capacity both to increase incubation effort and brood care investment.     

Are incubation costs in female pied flycatchers expressed in humoral immune responsiveness or breeding success?

Although clutch size variation has been a key target for studies of avian life history theory, most empirical work has only focused on the ability of parents to raise their altricial young. In this study, we test the hypothesis that costs incurred during incubation may be an additional factor constraining clutch size in altricial birds. In the pied flycatcher (Ficedula hypoleuca), we manipulated the incubation effort of the female by enlarging and reducing clutch sizes. To manipulate incubation effort only, the original clutch sizes were restored shortly after hatching. We found that fledging success was lower among broods whose clutches were enlarged during incubation. There was, however, no effect of manipulation on female body condition or on their ability to mount a humoral immune response to diphtheria or tetanus toxoid during the incubation or nestling provisioning period. Instead, we found that the original clutch size was related to the immune response so that females with seven eggs had significantly lower primary antibody responses against tetanus compared to those with six eggs. Our results suggest that incubating females are not willing to jeopardise their own condition and immune function, but instead pay the costs of incubating a larger clutch by lower offspring production. The results support the view that costs of producing and incubating eggs may be substantial and hence that these costs are likely to contribute to shaping the optimal clutch size in altricial birds.     

Does the cost of incubation set limits to clutch size in common eiders Somateria mollissima?

We examined the effect of natural clutch size on the cost of incubation in a population of common eiders Somateria mollissima nesting in Tromsø, northern Norway. The body condition of females at day 5 in the incubation period was not related to clutch size (3–6 eggs), but females incubating large clutches lost more mass and had a lower body condition at day 20 in the incubation period than females incubating small clutches. Females incubating large clutches had a slightly shorter incubation period and a lower egg predation rate. The results do not support the hypothesis that the female's ability to produce eggs is the only ultimate control of clutch size in eider. Instead the results suggest that there may be an interaction between the allocation of body reserves to eggs and incubation, and that females producing large clutches allocate more of their body reserves to incubation than females producing small clutches, in order to shorten the incubation period and to minimise the risk of predation on eggs.     

The role of partial incubation and egg repositioning within the clutch in hatching asynchrony and subsequent effects on breeding success

The main mechanism to achieve hatching asynchrony (HA) for incubating birds is to start heating the eggs before clutch completion. This might be achieved through partial incubation and/or early incubation. Even in the absence of incubation behaviour during the laying phase, clutches still experience a certain degree of asynchrony. Recent studies have shown that eggs located in the centre of the nest receive more heat than peripheral ones during incubation. As eggs receiving more heat would develop faster, we hypothesized that HA should be shorter in nests where eggs were moved homogeneously along the centre–periphery space during incubation than in those nests where eggs repeatedly remained in the same locations, either centrally or peripherally. We explored the relative roles of egg repositioning and partial incubation in determining HA in wild birds by (1) removing eggs from 20 Great Tit Parus major nests on the day of laying and replacing them with fake eggs to avoid partial incubation, and returning them when full incubation began; (2) monitoring twice a day the position of each individually marked egg relative to the clutch centre during incubation, and estimating the coefficient of variation of the distances; and (3) determining HA in each nest. Preventing partial incubation reduced HA by 51% days in experimental nests. It also caused negative effects for the incubating females (lengthening the full incubation period) and positive effects for the brood (increasing fledging success). However, our hypothesis about the role of egg repositioning on HA was not supported: all the females moved the eggs with remarkable consistency, generally attaining a coefficient of variation of the distances around 33%, and it was not related to the HA experienced. We therefore conclude that partial incubation is an important factor regulating HA, and females compensate for the potential effects of differential heating by moving the eggs homogeneously within the clutch.     

Brood conspicuousness and clutch viability in male‐caring assassin bugs (Rhinocoris tristis)

Abstract 1. Conspicuousness to mates can bring benefits to both males (increased mating success) and females (reduced search costs), but also brings costs (e.g. increased predation and parasitism). Assassin bugs, Rhinocoris tristis, lay egg clutches either on exposed stems or hidden under leaves. Males guard eggs against parasitoids. Guarding males are attractive to females who add subsequent clutches to the brood. This is an excellent opportunity to study the effects of conspicuousness on the fitness of males and females. 2. Using viable eggs in a multi‐clutch brood as a correlate of fitness, the present study examined whether laying eggs on stems affected (1) female fitness, through exposure to parasitism and cannibalism, and (2) male fitness, through attracting further females. 3. Stem broods were more parasitised. However, males on stems accumulated more mates and more eggs, a net benefit even accounting for parasitism. The eggs gained from being on a stem were cannibalised. By contrast, higher mortality on stems suggests that females should gain by ovipositing on leaves. To the extent that egg viability represents fitness, male and female interests may therefore differ. This suggests a potential for sexual conflict that may affect other species with male care. 4. Despite higher costs, females actually initiated more broods, and subsequently added bigger clutches to broods, on stems than under leaves. This suggests either that viable eggs do not reflect fitness, or that females laid in unfavourable locations. The key is now to address lifetime fitness, since unmeasured factors may affect offspring viability post‐hatching, and to investigate who controls the location of oviposition in R. tristis.     

Cascading costs of reproduction in female house wrens induced to lay larger clutches

In many species, females produce fewer offspring than they are capable of rearing, possibly because increases in current reproductive effort come at the expense of a female's own survival and future reproduction. To test this, we induced female house wrens (Troglodytes aedon) to lay more eggs than they normally would and assessed the potential costs of increasing cumulative investment in the three main components of the avian breeding cycle – egg laying, incubation and nestling provisioning. Females with increased clutch sizes reared more offspring in the first brood than controls, but fledged a lower proportion of nestlings. Moreover, nestlings of experimental females were lighter than those of control females as brood size and prefledging mass were negatively correlated. In second broods of the season, when females were not manipulated, experimental females laid the same number of eggs as controls, but experienced an intraseasonal cost through reduced hatchling survival and a lower number of young fledged. Offspring of control and experimental females were equally likely to recruit to the breeding population, although control females produced more recruits per egg laid. The reproductive success of recruits from broods of experimental and control females did not differ. The manipulation also induced interseasonal costs to future reproduction, as experimental females had lower fecundity than controls when breeding at least 2 years after having their reproductive effort experimentally increased. Finally, females producing the modal clutch size of seven eggs in their first broods had the highest lifetime number of fledglings.     

Costs of incubation and immunocompetence in the collared flycatcher

This paper investigates the costs of incubation in terms of reduced reproductive success and investigates whether incubation competes with immune function for resources. I performed a clutch size manipulation experiment in which two eggs were either removed from or added to the nests of collared flycatchers, Ficedula albicollis, for 1 week during incubation and subsequently returned to their original nests before hatching. To induce immune response, the females were challenged with sheep red blood cells. While the duration of incubation, hatching success and fledgling number did not differ between experimental groups, fledgling condition was significantly lower in broods that had been enlarged during incubation. Neither the females' condition nor their ability to respond to a novel antigen differed between treatments. The relationship between antibody production and female condition was significantly positive, but only among females incubating reduced clutches. I conclude that the costs of incubation in the collared flycatcher are not negligible and are manifested only at the chick-rearing phase.     

How avian incubation behaviour influences egg surface temperatures: relationships with egg position,development and clutch size

While understanding heat exchange between incubating adults and their eggs is central to the study of avian incubation energetics, current theory based on thermal measurements from dummy eggs reveals little about the mechanisms of this heat exchange or behavioural implications for the incubating bird. For example, we know little about how birds distribute their eggs based on temperature differences among egg positions within the nest cup. We studied the great tit Parus major, a species with a large clutch size, to investigate surface cooling rates of individual eggs within the nest cup across a range of ambient temperatures in a field situation. Using state‐of‐the‐art portable infrared imaging and digital photography we tested for associations between egg surface temperature (and rate of cooling) and a combination of egg specific (mass, shape, laying order, position within clutch) and incubation specific (clutch size, ambient temperature, day of incubation) variables. Egg surface temperature and cooling rates were related to the position of the eggs within the nest cup, with outer eggs being initially colder and cooling quicker than central eggs. Between foraging bouts, females moved outer eggs significantly more than centrally positioned eggs. Our results demonstrate that females are capable of responding to individual egg temperature by moving eggs around the nest cup, and that the energy cost to the female may increase as incubation proceeds. In addition, our results showing that smaller clutches experience lower initial incubation temperatures and cool quicker than larger clutches warrant further attention for optimal clutch size theory and studies of energetic constraints during incubation. Finally, researchers using dummy eggs to record egg temperature have ignored important elements of contact‐incubation, namely the complexity of how eggs cool and how females respond to these changes.     

The significance of clutch overlap in Great Tits Parus major

We studied the effects of manipulation of the size of first broods in the Great Tit Parus major on the size and breeding success of second clutches and its relation to the degree of clutch overlap. The rearing of first brood fledglings always overlapped with the laying of the second clutch and in most cases also with the incubation period of the latter. The degree of clutch overlap depended on the size of the first brood, being less when the first brood was large. Clutch overlap also increased with season. Mechanisms affecting the timing of laying of second clutches are discussed. A large first brood imposed reproductive costs. It affected the size of the second clutch by causing it to be delayed; second clutch size decreases with season. It affected the post-fledging survival of second brood young as, in this population, this decreases with fledging date. The breeding success of second clutches was, however, not affected by the size of the first brood, but instead by the weight of the female when rearing the first brood.     

Incubation capacity limits maximum clutch size in black-tailed gulls Larus crassirostris

Factors determining clutch size of birds have long been the central issue in studies in life histories. It is assumed that the configuration of brood patches could limit the maximum clutch size. To test this hypothesis we manipulated clutch sizes and measured egg temperature as well as reproductive consequences in black-tailed gulls Larus crassirostris , which usually lay two egg clutches and have three brood patches. Mean egg temperature in 4-egg clutches (32.6±1.0°C) was significantly lower than in 2-egg (34.6±0.4°C) and 3-egg clutches (34.1±0.4°C), because egg temperature of the coolest egg within a 4-egg clutch was often substantially lower than the other three eggs. The proportion of eggs hatching from 4-egg clutches (11.6%) was lower than those of 2-egg (49.1%) and 3-egg clutches (52.0%). Four-egg clutches had longer incubation periods (29.6±1.3 day) than 2-egg (28.1±1.7 day) and 3-egg clutches (28.0 ±1.3 day). The results indicate that incubation capacity, which may be determined by the configuration of brood patches, limits the maximum clutch size in black tailed gulls, but not the actual clutch size typically laid.     

Costs and response to conspecific brood parasitism by colonial red-breasted mergansers

Costs of conspecific brood parasitism (CBP) are expected to be influenced by a species’ life history traits. Precocial birds lay large clutches, and clutches that have been enlarged by CBP can affect host fitness through a longer incubation period, displaced eggs, and lower hatching success. We examined costs and response to CBP by hosts in a population of colonial red-breasted mergansers (Mergus serrator; n?=?400 nests over 8 years) within which 29% of parasitized clutches were enlarged considerably (≥?15 eggs). Length of the incubation period did not increase with clutch size. The mean number of eggs displaced from a parasitized nest during incubation (2.8) was 2×?greater than at an unparasitized nest (1.4). Hatching success declined by 2% for each additional egg in the nest. Thus, for a nest with?≥?15 eggs, one or more fewer host eggs hatch relative to an unparasitized nest with the same number of host eggs, assuming equal probability of success for all eggs. Hosts were 40% more likely to desert nests receiving 2 or 6 experimental eggs relative to unparasitized control nests, although it is unknown whether hens deserting a nest renested elsewhere. Our study indicates that costs of CBP to hosts during nesting may be limited to those red-breasted mergansers incubating the largest clutches (≥?15 eggs), and it raises questions about the adaptive significance of deserting a parasitized clutch.     

Clutch size manipulation, hatching success and offspring phenotype in the ball python (Python regius)

In a diverse array of avian and mammalian species, experimental manipulations of clutch size have tested the hypothesis that natural selection should adjust numbers of neonates produced so as to maximize the number of viable offspring at the end of the period of parental care. Reptiles have not been studied in this respect, probably because they rarely display parental care. However, females of all python species brood their eggs until hatching, but they do not care for their neonates. This feature provides a straightforward way to experimentally increase or reduce clutch size to see whether the mean clutch size observed in nature does indeed maximize hatching success and/or optimize offspring phenotypes. Eggs were removed or added to newly laid clutches of Ball Pythons ( Python regius ) in tropical Africa (nine control clutches, eight with 50% more eggs added, six with 42% of eggs removed). All clutches were brooded by females throughout the 2-month incubation period. Experimental manipulation of clutch-size did not significantly affect the phenotypes (morphology, locomotor ability) of hatchlings, but eggs in 'enlarged' clutches hatched later, and embryos were more likely to die before hatching. This mortality was due to desiccation of the eggs, with females being unable to cover 'enlarged' clutches sufficiently to retard water loss. Our results support the notion of an optimal clutch size, driven by limitations on parental ability to care for the offspring. However, the proximate mechanisms that generate this optimum value differ from those previously described in other kinds of animals. © 2003 The Linnean Society of London . Biological Journal of the Linnean Society 2003, 78 , 263–272.     

Age-dependent reproductive costs and the role of breeding skills in the Collared flycatcher

This study addressed whether there are any age‐related differences in reproductive costs. Of especial interest was whether young individuals increased their reproductive effort, and thereby their reproductive cost, as much as older birds when brood size was enlarged. To address these questions, a brood‐size manipulation experiment with reciprocal cross‐fostering of nestlings of young and middle‐aged female Collared flycatchers, Ficedula albicollis, was performed on the Swedish island of Gotland. Nestlings’ body mass, tarsus length and survival were recorded to estimate the parental ability and parental effort of the experimental female birds. Female survival and clutch size were recorded in the following years to estimate reproductive costs. We found that middle‐aged female flycatchers coped better with enlarged broods than younger females or invested more in reproduction. In the following year, young female birds that had raised enlarged broods laid smaller clutches than the females from all the other experimental groups. This result shows that the young female birds pay higher reproductive costs than the middle‐aged females. Both young and middle‐aged female flycatchers seemed to increase their reproductive effort when brood size was increased. However, such an increase resulted in higher reproductive costs for the young females. The difference in reproductive costs between birds of different ages is most likely a result of insufficient breeding skills of the young individuals.     

ESTABLISHMENT OF WEIGHT HIERARCHIES IN THE BROODS OF HOUSE MARTINS DELICHON URBICA

Nestling birds may differ in size and weight on the first day a clutch is fully hatched, mainly because eggs within clutches hatch over a period of several days. This asynchronous pattern of hatching is usually thought to facilitate brood reduction when the food supply is unpredictably restricted. The purpose of the study reported here was to examine the contribution of egg-weight, clutch-size, hatching spread, food supply and season to weight differences in newly hatched broods of the House Martin. At laying, heavy eggs had a greater moisture and dry weight content than light eggs and immediately before hatching there was a correlation between initial egg-weight and the dry weight of embryo and yolk. Heavier clutches also tended to give rise to heavier hatchlings. There was, however, no correlation of fresh egg-weight with the dry weight of embryos alone and the relative dry weight of embryos in a clutch was dependent on laying sequence. Hatching spread (the number of days between the emergence from the egg of the first and the last hatchling of the clutch) was 0.75 ± 0.46 days for clutches of two and increased with the size of the clutch up to 1.80 ± 0.79 days for clutches of five. When food was scarce during laying, hatching spread was greater. Weight difference in newly hatched broods was correlated with hatching spread and moreover in multivariate analysis was also correlated with periods of food scarcity during laying. It was concluded that all examples of weight hierarchies among hatchlings should not be considered adaptive; in some cases they may be imposed by food scarcity. This can lead to mortality of the runs even if food is plentiful. When the weight hierarchy is not adversely accentuated by food scarcity it may function as previously suggested, to allow brood reduction. Alternatively, particularly among House Martins, it may spread out the peak food needs of individual nestlings thereby spreading the demand on the adults.     

Male Isaza (Gymnogobius isaza, Gobiidae) prefer large mates: a counterstrategy against brood parasitism by conspecific females

Like many other gobies, male Isaza (Gymnogobius isaza) which are endemic to Lake Biwa, Japan, exclusively care for broods in nests. This goby may have an optimal range of brood size (i.e., an average clutch size of about 2000–3000 eggs) within which they may produce larger numbers of hatching young because much larger broods may be destroyed by fungal infection before hatching. This optimal brood size hypothesis (Takahashi et al. in J Ethol 22:153–159, 2004) predicts that (1) after spawning, both males and females will refuse additional spawning by other gravid females (second females) to keep brood sizes within optimal ranges, (2) larger fish will repel second females more successfully than will smaller fish, and thus, (3) both sexes prefer larger mates. To examine these predictions, we first observed Isaza’s aggressive behaviors in aquaria and investigated whether fish attacked and repelled second females that were introduced after spawning, and, if so, what were the sizes of fish that did so. Large fish, regardless of sex, aggressively prevented second females from entering the nest, but second females larger than the pairs displaced the pair females forcibly and spawned eggs into their clutches. Mate choice experiments showed that males preferred large females. Although females’ choice of large mates was not confirmed, many results may largely coincide with the predictions of the optimal brood size hypothesis. Thus, Isaza males’ choice of large mates will be advantageous for defending against brood parasitism by conspecific females and for achieving optimal clutch size.     

Incubation capacity contributes to constraints on maximal clutch size in Brent Geese Branta bernicla nigricans

Lack ( 1967 ) proposed that clutch size in species with precocial young was determined by nutrients available to females at the time of egg formation; since then others have suggested that regulation of clutch size in these species may be more complex. We tested whether incubation limitation contributes to ultimate constraints on maximal clutch size in Black Brent Geese (Black Brant) Branta bernicla nigricans. Specifically, we investigated the relationship between clutch size and duration of the nesting period (i.e. days between nest initiation and the first pipped egg) and the number of goslings leaving the nest. We used experimental clutch manipulations to assess these questions because they allowed us to create clutches that were larger than the typical maximum of five eggs in this species. We found that the per‐capita probability of egg success (i.e. the probability an egg hatched and the gosling left the nest) declined from 0.81 for two‐egg clutches to 0.50 for seven‐egg clutches. As a result of declining egg success, clutches containing more than five eggs produced, at best, only marginally more offspring. Manipulating clutch size at the beginning of incubation had no effect on the duration of the nesting period, but the nesting period increased with the number of eggs a female laid naturally prior to manipulation, from 25.4 days (95% CI 25.1–25.7) for three‐egg clutches to 27.7 days (95% CI 27.3–28.1) for six‐egg clutches. This delay in hatching may result in reduced gosling growth rates due to declining forage quality during the brood rearing period. Our results suggest that the strong right truncation of Brent clutches, which results in few clutches greater than five, is partially explained by the declining incubation capacity of females as clutch size increases and a delay in hatching with each additional egg laid. As a result, females laying clutches with more than five eggs would typically gain little fitness benefit above that associated with a five‐egg clutch.     

ON MULTIPLE BROODS AND THE BREEDING STRATEGY OF ARCTIC SANDERLINGS

Sanderlings on Bathurst Island in the Canadian arctic have two patterns of incubation. At some nests the eggs are covered soon after the fourth egg has been laid and at others incubation is delayed for 5–6 days. Because the delay is about the same time required to lay a second clutch and because a single individual alone incubates at any one nest, we suspected that Sanderlings might normally lay two clutches in a season, the male caring for one brood and the female for the other.
Histological and gross examination of the ovaries of two females taken as the birds began incubation showed eight freshly ovulated follicles in each female. The size gradation and histological appearance of the follicles indicated that two clutches of four eggs each had been laid within 8–10 days by a single female. The ovary of one female had additional large yolky follicles, suggesting a physiological capability of further ovulations.
Field conditions in the arctic summer are highly variable, and the small eggs and the rapid sequence of broods of Sanderlings may be breeding adaptations that permit them to multiply the traditional wader clutch of four eggs by 2 or 3 in favourable years. Selection for mating systems characterised by brief pair bonds and by polyandry is expected in precocial birds where some broods are incubated and cared for by the male, but further field work is required to determine the precise mating system of Sanderlings.     

CLUTCH-SIZE, INCUBATION AND HATCHING SUCCESS IN THE HOUSE SPARROW AND TREE SPARROW PASSER SPP. AT OXFORD

Clutch-size, incubation and hatching success were studied in P. domesticus and P. montanus in 1961 and 1963–64 at Oxford. The most frequent clutch-size was four eggs in P. domesticus and five eggs in P. montanus . With one exception, colonies of P. domesticus showed no significant annual or local variations in its mean clutch-size; in P. montanus , however, there were significant annual variations in the mean clutch-size. Both species showed a seasonal increase followed by a decrease in their mean clutch-sizes.
Partial incubation occurred during the laying period of the clutch; sufficient incubation for continuous development of the embryo was apparently achieved when the last egg had been laid in clutches of two and three eggs in P. domesticus and in clutches of four eggs in P. montanus , but when the penultimate egg had been laid in larger clutches of both species. On average, hatching in P. domesticus occurred more or less synchronously in all eggs in clutches of two and three eggs, and in all eggs except the last one laid in larger clutches; the last egg in the larger clutches hatched up to a day after the others. It is suggested that this pattern of hatching was brought about by the pattern of incubation during the laying period.
P. domesticus had a lower hatching success than P. montanus , probably because fewer of its eggs were fertile.     

What limits clutch size? A test of the incubation‐capacity hypothesis in a high‐elevation population of Mountain Bluebirds

Most studies of factors that limit the number of eggs that birds lay have focused on the disadvantages of having too many young to feed. Less attention has been paid to the consequences of having a large number of eggs to incubate. The incubation‐capacity hypothesis proposes that females lay as many eggs as they can effectively incubate. We tested this hypothesis in 2018 in a montane population of Mountain Bluebirds (Sialia currucoides). Most females in this population lay five or six eggs; clutches of seven occur, but are rare. We added eggs to some nests, forcing females to incubate seven eggs, while leaving other nests as controls. Among females completing incubation, those with enlarged clutches hatched as many eggs as did control females, and did so in the same amount of time. This was despite an extended period of unusually cold and often wet weather that occurred when many females were incubating. Our results firmly reject the suggestion that females typically lay no more than six eggs because they cannot effectively heat seven eggs. One or more other factors must limit clutch size. One possible factor is suggested by the fact that during the period of inclement weather, more females with enlarged clutches than control females appeared to abandon nests before completing incubation. Because larger clutches require more energy to incubate, females with seven eggs during energetically stressful conditions could more quickly reach the point where they lack sufficient energy for both incubation and self‐maintenance. Such conditions may occur frequently enough in the montane environment that, on average, laying seven eggs results in reduced lifetime reproductive success.     

Brood parasitism,relatedness and sociality: a kinship role in female reproductive tactics

Conspecific brood parasitism (CBP) is a reproductive tactic in which parasitic females lay eggs in nests of other females of the same species that then raise the joint brood. Parasites benefit by increased reproduction, without costs of parental care for the parasitic eggs. CBP occurs in many egg‐laying animals, among birds most often in species with large clutches and self‐feeding young: two major factors facilitating successful parasitism. CBP is particularly common in waterfowl (Anatidae), a group with female‐biased natal philopatry and locally related females. Theory suggests that relatedness between host and parasite can lead to inclusive fitness benefits for both, but if host costs are high, parasites should instead target unrelated females. Pairwise relatedness (r) in host–parasite (h‐p) pairs of females has been estimated using molecular genetic methods in seven waterfowl (10 studies). In many h‐p pairs, the two females were unrelated (with low r, near the local population mean). However, close relatives (r = 0.5) were over‐represented in h‐p pairs, which in all 10 studies had higher mean relatedness than other females. In one species where this was studied, h‐p relatedness was higher than between nesting close neighbours, and hosts parasitized by non‐relatives aggressively rejected other females. In another species, birth nest‐mates (mother–daughters, sisters) associated in the breeding area as adults, and became h‐p pairs more often than expected by chance. These and other results point to recognition of birth nest‐mates and perhaps other close relatives. For small to medium host clutch sizes, addition of a few parasitic eggs need not reduce host offspring success. Estimates in two species suggest that hosts can then gain inclusive fitness if parasitized by relatives. Other evidence of female cooperation is incubation by old eider Somateria mollissima females of clutches laid by their relatives, and merging and joint care of broods of young. Merging females tended to be more closely related. Eiders associate with kin in many situations, and in some geese and swans, related females may associate over many years. Recent genetic evidence shows that also New World quails (Odontophoridae) have female‐biased natal philopatry, CBP and brood merging, inviting further study and comparison with waterfowl. Kin‐related parasitism also occurs in some insects, with revealing parallels and differences compared to birds. In hemipteran bugs, receiving extra eggs is beneficial for hosts by diluting offspring predation. In eggplant lace bugs Gargaphia solani, host and parasite are closely related, and kin selection favours egg donation to related females. Further studies of kinship in CBP, brood merging and other contexts can test if some of these species are socially more advanced than presently known.