Temporal Variation in Decisions about Parental Care in Bluegill, Lepomis macrochirus

Parental investment theory states that an individual will trade‐off present and future reproductive potential to maximize lifetime reproductive success. Only when parental care is costly in terms of reduced future reproductive potential should individuals be sensitive to changes in the value of current offspring and adjust their care. Here, we examine temporal variation in parental care decision‐making in bluegill (Lepomis macrochirus), in which care is provided by males called ‘parentals’. Previous research has shown that parentals that nest early in the breeding season are in higher energetic condition than those that nest later, and early nesting males appear not to pay an opportunity cost to their care in terms of reduced future reproductive potential. Early nesting males also may have higher paternity in their broods than later nesting males. To examine the parental care decisions made by early and mid‐season nesting parentals, we experimentally reduced males’ perceived paternity by swapping eggs between nests. We found that experimental males that nested early in the breeding season adjusted their brood defence behaviour similarly to control males, which had sham egg swaps performed. Conversely, experimental males that nested mid‐season significantly decreased their brood defence behaviour after the manipulation as compared with control males. Thus, unlike mid‐season nesting males, early nesting males appear relatively insensitive to changes in brood value (paternity), possibly because early nesting males pay little cost in terms of reduced future reproductive potential to providing full care or because these males have a predisposition to high paternity.     

Contrasting breeding strategies of two sympatric orioles: first documentation of double brooding by Orchard Orioles

ABSTRACT.   Although many studies have revealed intraspecific variation in the number of broods raised per breeding season, less is known about the causes of variation between species. We monitored the breeding behavior of congeners traditionally thought to raise one brood per breeding season, Orchard Orioles ( Icterus spurius ) and Baltimore Orioles ( I. galbula ). Both species have similar habitat and nesting requirements. We captured and banded orioles at three study sites in Maryland in 2004 and 2005. No pairs of Baltimore Orioles ( N = 26) were double brooded, whereas 8 of 24 pairs (33%) of Orchard Orioles had a second brood and 7 of 16 pairs (44%) of Orchard Orioles that successfully raised a first brood also successfully raised a second brood during the same breeding season. Our findings provide the first documentation of double brooding by Orchard Orioles. The timing of molt and the amount of time and energy devoted to postfledgling parental care may contribute to this difference between species in terms of the number of broods per breeding season.     

Polygyny in the tree swallow Tachycineta bicolor: a result of the cost of searching for an unmated male

The costs imposed by parental care duties on an individual's future survival and reproduction generate conflicts because parents should attempt to minimize their investment in the present brood, and exploit the parental care of the other parent. This conflict is likely to contribute to cases of both polygamy and desertion. Here, we study the costs of polygyny in the tree swallow Tachycineta bicolor , using observations on 52 nests that were attended by polygynous males over 14 y. Both females mated to polygynous males paid reproductive costs at several stages of the nesting cycle. Clutches laid by social mates of polygynous versus monogamous males did not differ in size. However, initial brood sizes for polygynously mated females were lower because a higher proportion of their eggs failed to hatch. Likewise, fledging success was lower and nest predation rates were higher, perhaps reflecting the direct or indirect effects of reduced male attention. These results demonstrate that females pay a productivity cost when breeding with reduced male parental care. In contrast, polygynous males fledge on average more young than monogamous ones and clearly benefit from the association. We suggest that a mate-search cost is leading to the few cases of polygamous males: although females are likely evaluating males for their prospective dedication to the breeding attempt, in a short-lived bird with a short breeding season, the cost to females of searching for a more dedicated male is the risk of not breeding at all.     

Strict monogamy in a semi-colonial passerine: the Jackdaw Corvus monedula

In this study we used DNA fingerprinting to provide an accurate measure of paternity in a nest-box colony of Jackdaws. The species is a semi-colonial, socially monogamous passerine which establishes long-term pair bonds. Parents raise a single annual brood in which 50% brood reduction is commonplace. However, nest sites are a limited resource, non-breeding adults are also common around colonies and males are frequently separated from their incubating females during the fertile egg-laying period so that opportunities arise for extra-pair copulation. The parentage analysis, however, revealed no cases of extra-pair fertilisation (EPF) or intra-specific brood parasitism. Therefore fledgling output or survival is likely to be a good measure of individual reproductive success in this species. The lack of EPFs is not explained by nesting synchrony and we discuss the relative costs and benefits to females of seeking EPFs; the likelihood that paternal care and a life-history strategy similar to many long-lived non-passerines may also constrain the species to monogamy.     

Successive clutches and parental roles in waders: the importance of timing in multiple clutch systems

Production of successive clutches within the same breeding season has received less attention than many other aspects of avian reproduction. Waders are of particular interest because in these birds, multiple clutches are associated with at least three different breeding systems: double-clutching (uniparental care), monogamous double-brooding (biparental care) and polyandry (uni- or biparental care). Data from eight species and twelve breeding populations suggest that early second clutches, and thus brood overlap, are associated with parental role division and uniparental care, whereas species or populations with biparental care tend to have long intervals between successive clutches. We suggest that ecological factors influencing the relative timing of the second clutch will have consequences for the parental care system. In particular, conditions that favour early laying of the second clutch (large brood overlap) are likely to lead to parental role division, as found in double-clutching species. Factors determining the timing of second clutches are discussed, as are possibilities for testing these ideas.     

Sexual aggression by intruders in hooded crow <Emphasis Type="Italic">Corvus cornix</Emphasis>

The hooded crow Corvus cornix is a west Palaearctic, solitary nesting, monogamous corvid. In the breeding season, populations are characterized by a social organization wherein breeding pairs are territorial and non-breeding individuals, called floaters, live in flocks. During a study of the breeding ecology of the hooded crow, conducted in a protected flooded area, we monitored nests with video cameras. We recorded two separate incidents when intruders attacked a female at the nest. We believe that she remained in the nest in order to prevent the strangers cannibalizing the nestlings by mantling over the brood. The spatio-temporal occurrence of these attacks suggests that the observed behaviour is intraspecific sexual aggression wherein non-breeding males mounted an immobilized female.     

An experimental test on time constraint and sexual conflict over parental care

Because parental care is costly, a sexual conflict between parents over parental investment is expected to arise. Parental care behavior is an adaptive decision, involving trade‐offs between remating, and consequently desertion of the brood, and continuing parental effort. If the main advantage of desertion is remating, then this will be a time constraint, because the deserting individual will require a certain minimum period of time to breed again in the same breeding season. So, a short breeding season should force certain individuals to desert the first brood to have enough time to successfully complete their second breeding attempt. The rock sparrow, Petronia petronia, is an unusual species in which brood desertion can occur in both sexes and the breeding season is quite short so it is a good species to investigate the role of time constraint on brood desertion. For 3 years, I investigated the brood desertion modality of the rock sparrow. Then, for 2 years, I removed a group of experimental nest boxes during the autumn. Later, I re‐installed the experimental nest boxes after the start of the breeding season (2 weeks after the first egg was laid), mimicking a shortening of the breeding season for the (experimental) pairs that used experimental nest boxes. I found that in the experimental pairs, the percentage of deserting individuals was significantly higher than in the control groups, and the deserting individuals were older females. This experiment adds to our knowledge of timing of reproduction effects on individual decisions to desert by showing that a short and delayed breeding season may have different effects on males and females. To my knowledge, this is the first experimental study that demonstrates a direct link between time constraint and brood desertion.     

Plasma levels of androgens and cortisol in relation to breeding behavior in parental male bluegill sunfish, Lepomis macrochirus

Like many teleosts, male bluegill sunfish (Lepomis macrochirus) provide sole parental care. To understand some of the proximate costs of parental care, we measured body condition and plasma levels of testosterone (T), 11-ketotestosterone (11KT) and cortisol in nesting bluegill males during pre-spawning, spawning and parental care stages. T and 11KT were at their highest mean levels during the pre-spawning period and decreased to lower levels early during the parental care period before rising again when the eggs hatched. Cortisol levels fluctuated across the breeding stages, but there was a noticeable increase from low levels on the day of spawning during the first 2 days of parental care when egg fanning is most intense. Levels of all hormones varied considerably among males, with androgen levels often correlating positively with a male's body condition. We also demonstrate, using a brood reduction experiment and repeated sampling of known individuals, that the presence of eggs affects hormone levels shortly after eggs hatch. Parental males in better body condition had higher levels of androgens during parental care. Males that were known to renest later in the season also had higher androgen levels and were in better body condition during the first nesting bout than males only known to have nested once. However, circulating levels of cortisol did not differ significantly between these groups. We discuss our findings in the context of proximate and ultimate costs of parental care and propose several reasons why elevated androgen levels may not be as incompatible with the expression of paternal care in male teleosts, as compared with avian and mammalian fathers.     

Sex roles in the monogamous Purple Sandpiper Calidris maritima in Svalbard

The Purple Sandpiper Calidris maritima, an Arctic shorebird, displays unusual sex roles during breeding. In a 5-year study in Svalbard, in the Norwegian high Arctic, data were collected on the roles of the sexes in this species. Purple Sandpipers were similar to most shorebirds in that males actively courted females, established territories and vigorously defended their territories from intruders. Both sexes shared incubation duties approximately equally over the entire 21-day incubation period. Males incubated very little initially, but increased their effort significantly during the first 11 days of incubation to over 50% of the time from days 11–21. However, Purple Sandpipers contrasted with most other shorebird species in that females discontinued their breeding efforts at hatching. In nearly all cases, broods were attended solely by the male until the chicks reached fledging age (or even longer). Nonetheless, the pair bond must be described as monogamous because neither males nor females were found to re-mate or lay a second clutch during a season. This pattern of parental care, which is found only in very few other shorebird species, is discussed in an evolutionary context.     

Temporal patterns of circulating LH and steroid hormones in a brood parasite, the Brown-headed cowbird, Molothrus ater.

The reproductive cycle of monogamous passerines is characterized by a sequence of events leading from courtship and pair formation through nest building, egg laying and incubation, and ending with the care of young. Each of these stages is accompanied by equally predictable changes in the pattern of hormone secretion. This report describes endocrine changes in the plasma of free-living Brown-headed cowbirds, Molothrus uter , during a breeding season. Changes in some morphological parameters are also presented.
Cowbirds are brood parasites that show no parental behaviour and lay eggs almost daily for approximately eight weeks. Males are not territorial, but they mate guard extensively. Hcncc, their breeding biology is quite different from that of the nesting species described above. They are hormonally distinct, as well. Luteinizing hormone rises in the spring and remains elevated throughout May and most of June. By July, when breeding has ceased, plasma luteinizing hormone levels are at pre-breeding values. Plasma androgens are also maintained at peak levels for an extended period of time, although they decline briefly between the establishment of dominance hierarchies and the onset of breeding. Circulating levels of corticosterone are elevated until the middle of the breeding season, when they begin to fall, although here, too, there is a transient depression just prior to breeding. In the light of the studies on nesting species, these data suggest that hormone profiles are closely correlated with the breeding and social activities that characterize the species under investigation.     

Food availability and the male's role in parental care in double-brooded Treecreepers Certhia familiaris

The aim of this work was to examine differences in paternal and maternal care in a double-brooded, monogamous species, the Treecreeper Certhia familiaris, in relation to food availability. As a measure of parental care, we recorded the hourly feeding activity of parents when the nestlings from their first and second breeding attempts were 7 and 12 days old. Feeding frequency of the first brood increased with the age of the nestlings and also with the brood size when 12 days old. While the feeding activities of the females were similar with respect to the first and second broods, the males were less active and failed to provide any food to their nestlings in 15 cases out of 28 second broods. In spite of this, the fledglings from the second broods were heavier than those in the first. Such a pattern of male behaviour was possible without being a disadvantage to the chicks because the food supply increased during the breeding season and the female could provide food for the young alone. Thus paternal care was particularly important in times of poor food supply, i.e. during the first brood, where the extent of these males' activity in feeding the 7-day-old nestlings was positively correlated with the average mass of the nestlings. Our results support the idea that the male of monogamous, altricial bird species often makes important contributions to raising the young, especially during periods when it is difficult for the female to do so alone. Males show flexibility in their pattern of parental care, and male Treecreepers change their contribution to the first and second broods within the same season.     

Serial monogamy in the European long-snouted seahorse Hippocampus guttulatus

Seahorses (Hippocampus spp.) are non-sex-role-reversed members of the Syngnathidae family that provide extensive brood care. Previous studies of seahorses have revealed monogamy within a single brood, but their longer term mating system had not been comprehensively evaluated. The parental contribution to 29 wild-born broods of Hippocampus guttulatus, sampled from six Portuguese populations with differing seahorse densities and sex ratios, was assessed using microsatellite DNA markers. To assess the longer term genetic mating system of this species parentage was determined in eleven broods sampled from a captive population over two breeding seasons. Genetic data suggest that this socially polygamous seahorse is serially monogamous across breeding seasons, i.e. monogamous within a season but may switch mates between seasons, and that differing population densities and sex ratios do not affect the mating system.     

The effect of environmental factors on the nesting and courtship behaviour ofTilapia zillii in Lake Kinneret (Israel)

Tilapia zillii is a polyphilic, guarding nest spawner which is widespread in west and north Africa and in the Middle East. A study of the nesting and courtship behaviour of this species in Lake Kinneret (Sea of Galilee), revealed that nesting and brood care differ according to the exposure of the nesting site to wave action and the nature of the substrate. The nests are simple and parental care is abbreviated in exposed, sandy sites whereas elaborate nests with brood chambers are excavated in sheltered areas with clayey substrates, and parental care is extensive. The implications of these findings to studies on cichlid breeding and the conservation ofT. zillii in Lake Kinneret are discussed.     

SHOREBIRDS OF COASTAL BUENOS AIRES PROVINCE, ARGENTINA

The Pampas shorebird community is comprised of species from diverse taxonomic and faunal origin. Four major groups can be recognized: North American migrants residing in coastal Buenos Aires Province during their non-breeding season, Patagonian species wintering locally but breeding further south, Pampean species breeding locally, and Brazil–Paraguayan species present during their breeding season. Because of differences among these groups in the timing and migration patterns, community composition shows marked seasonal variation. Both nesting and non-breeding birds use a diversity of habitat types ranging from littoral sandy beaches to upland grasslands. Throughout the year, however, and in all habitat types, non-breeders predominate in species number and abundance. Intra- and inter-specific patterns of aggression suggest both that resources are limiting, and that non-breeding migrants may play an important role in determining breeding community composition.     

Effects of migration distance on life history strategies of Western and Semipalmated sandpipers in Perú

Migration distances of shorebird species correlate with life history strategies. To assess age‐specific migratory preparation and adult wing‐molt strategies, we studied Western Sandpipers (Calidris mauri) and Semipalmated Sandpipers (C. pusilla) with different migration routes at the Paracas National Reserve in Perú, one of the most austral non‐breeding areas for these sandpipers, from 2012 to 2015. Western Sandpipers breed near the Bering Sea, ~11,000 km from Paracas. Semipalmated Sandpiper populations at Paracas are a mixture of short‐billed birds from western Arctic breeding sites, plus long‐billed birds from eastern sites, ~8000 km distant. Adults of both species arrive in October with primary feathers already partially renewed so wing molt starts at sites further north. Semipalmated Sandpipers with longer bills completed wing molt later than shorter billed birds. Adults of both species prepared for migration in February and March. No juvenile Western Sandpipers prepared for migration, confirming the “slow” over‐summering life history strategy of more southerly non‐breeding populations. Juvenile Semipalmated Sandpipers showed bimodality in strategies. Most showed no migratory preparation, but, during three non‐breeding periods, from 27% to 31% fattened, molted, and partially replaced outer primaries during the pre‐migratory period. Juveniles with longer culmens were heavier and tended to have more alternate plumage. Juveniles that were partially molting primaries had longer culmens and more alternate plumage. Juvenile Semipalmated Sandpipers from eastern‐breeding populations thus have a higher propensity for a fast life history strategy, and western birds a slow one, at this non‐breeding site in Peru. Western‐breeding Semipalmated Sandpiper populations thus resemble Western Sandpipers, suggesting a common, possibly distance‐related, effect on life history strategy.     

Physiological costs and carry-over effects of avian interspecific brood parasitism influence reproductive tradeoffs

Although models of co-evolution between brood parasites and their hosts primarily focus upon the cost to hosts in the current reproductive bout, the impact of brood parasitism may carry over to future reproductive attempts by altering resource allocation. Glucocorticoid stress hormones help mediate resource allocation to reproduction, yet they have rarely been examined in brood parasitic systems. Here we determined if shifts in parental care and corticosterone had carry-over effects on future reproductive effort in the rufous-and-white wren (Thryophilus rufalbus), a host of the Central American striped cuckoo (Tapera naevia). We found that parasitized parents had significantly higher stress-induced, but not baseline, corticosterone than natural parents during the fledgling stage, which was associated with changes in parental care. The high investment in current reproduction while parasitized may be due to the value of fledged chicks in tropical systems. This maladaptive response by parasitized parents was associated with delayed re-nesting and a reduced likelihood of nesting in the subsequent breeding season. Although a reduction in future reproductive effort can result from a combination of factors, this work suggests that fitness costs of brood parasitism are mediated by changes in corticosterone and parental care behavior that carry over into subsequent breeding seasons.     

Mate desertion by male Great Reed Warblers Acrocephalus arundinaceus at the end of the breeding season

Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains. Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains. Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains. Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains. Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains.     

Parental care in Semipalmated Sandpipers Calidris pusilla: brood desertion by females

Although Semipalmated Sandpipers Calidris pusilla are monogamous, with biparental incubation, most females (86–97%) deserted their broods to the care of their mates, 0–11 days (average 6) after their eggs hatched. Males left the brood an average of 8 days later, shortly before or after the chicks fledged. In several instances, females that deserted in one year remained with the chicks the next year, and vice versa. Females deserted chicks at nests that hatched later in the season at an earlier age than those had hatched earlier ( r ²> o.6). Since females appeared to have an energy deficit in at least some years, and suffered higher mortality rates than males during breeding, it is possible that females deserted broods in order to take advantage of better feeding conditions at migratory stopovers in northeastern North America early in the season. There was little evidence of higher nesting success or earlier hatching date in reuniting pairs, although if both members of a pair returned to the breeding area, 80% reunited. Increased survival of their mate may be most advantageous to males in ensuring that they obtain a female the following year.     

Monogamous pair bonds and mate switching in the Western Australian seahorse Hippocampus subelongatus

Apparently monogamous animals often prove, upon genetic inspection, to mate polygamously. Seahorse males provide care in a brood pouch. An earlier genetic study of the Western Australian seahorse demonstrated that males mate with only one female for each particular brood. Here we investigate whether males remain monogamous in sequential pregnancies during a breeding season. In a natural population we tagged males and sampled young from two successive broods of 14 males. Microsatellite analyses of parentage revealed that eight males re‐mated with the same female, and six with a new female. Thus, in this first study to document long‐term genetic monogamy in a seahorse, we show that switches of mates still occur. Polygynous males moved greater distances between broods, and tended to have longer interbrood intervals, than monogamous males, suggesting substantial costs associated with the breaking of pair bonds which may explain the high degree of social monogamy in this fish genus.     

Brood provisioning rates and fledgling behavior of Cordilleran Flycatchers in southwestern Colorado

The behavior of young songbirds after fledging is one of the least understood phases of the breeding cycle, although parental provisioning rates and movement of fledglings are key to understanding life history evolution. We studied Cordilleran Flycatchers (Empidonax occidentalis) at two sites in southwestern Colorado, USA, from 2012 to 2017. We banded and sexed breeding adults to determine the relative contributions of males and females to nestling and fledgling care, and attached radio‐transmitters to nestlings to facilitate observations of brood behavior after fledging. Females made 60% and 78% of total observed feedings of nestlings and fledglings, respectively. Parental provisioning rates increased with nestling age, and per‐nestling provisioning rates increased with brood size. Parental provisioning rates declined just before fledging, then increased just after fledging. Fledging times of individuals in broods were asynchronous and concentrated during the late afternoon and early evening. Males stopped caring for fledglings before females even though this species is single‐brooded, with some late‐season broods being abandoned by males. Broods spent the first three weeks after fledging within 400 m of nests, after which they began to disperse. Most aspects of the breeding biology of Cordilleran Flycatchers in our study, including the duration of nestling and fledging periods, female‐dominated provisioning, and movement patterns of fledglings, were similar to those of other Empidonax species. However, the times when young fledged were not concentrated in the morning as reported in most other songbirds, and this result warrants additional study of the timing of fledging in ecologically and taxonomically similar species. The increased per‐nestling provisioning rate with increasing brood size was unexpected, and additional study is needed to determine if this increase results from a trade‐off between adult annual survival and productivity favoring increased provisioning of young in larger broods, or from the existence of high‐quality individuals where larger clutches and higher provisioning rates are linked.