Sex and individual differences in foraging behavior of Japanese cormorants in years of different prey availability

Japanese cormorants, Phalacrocorax capillatus, are sexually dimorphic seabirds with males that are heavier and that dive deeper than females. Sex differences in prey composition and foraging behavior of those rearing chicks at Teuri Island, Hokkaido, were examined by collecting food-loads from parents in 1992–1998 and by radio-tracking ten birds each in 1997 and 1998 when prey availability was different. Males fed more on benthic and epibenthic fishes (82% mass) than did females (34%) while females fed more on epipelagic and coastal fishes (53%) than did males (18%). Males made longer dives (53 s) at feeding sites closer to the island (7 km) than females (39 s, 13 km) in 1997. In 1998 when the availability of epipelagic fish seemed to be higher, there were no sex differences in dive duration and distance to the feeding sites (35 s and 9 km for males, 36 s and 10 km for females). This sex difference in foraging behavior with a poor availability of epipelagic fish suggests that high diving ability possibly enables males to feed on demersal fish. Birds specializing in coastal shallow waters around the island made long dives; hence they were probably foraging in bottom layers. Those foraging in deeper shelf waters made short dives and they were thought to forage in surface layers. Electronic Publication     

Population structure and spatial pattern in the dioecious shrub Ceratiola ericoides

Abstract. The dioecious shrub Ceratiola ericoides (Florida rosemary) dominates xeric, infrequently burned Florida scrub vegetation, often to the near-exclusion of other woody species. We studied the spatial pattern, age, sex and size structure of four populations in Florida, USA: two coastal scrub populations subject to recurrent local disturbances due to sand movement, and two inland scrub populations in sites periodically burned by stand-replacing fires. The age structure of individual genets was estimated from node counts and used to describe the age structure of the populations. The sex ratio of males to females was not significantly different from 1:1, except within a female-biased coastal population subject to frequent sand movement. Node counts indicated that the mean age for reproductive individuals was 15 - 16 yr for the inland populations and 13 - 16 yr for the coastal populations. In all sites, there was no difference in mean age between males and females. Vegetative reproduction was uncommon except for the least-disturbed coastal population where 72 % of the reproductive individuals originated through layering. Individuals were generally randomly dispersed at the coastal sites, whereas significant aggregation of males and females occurred in the inland sites where the populations were initiated following fire. Seedling recruitment was continuous in the disturbed coastal scrub site, where 35% of the individuals were juveniles. Most juveniles were dispersed from 0.5 to 0.75 m around females. At one of the inland sites, where juveniles comprised 11% of the population, juveniles were clustered at 0.25 to 5.75 m around females. Coastal populations were all-aged, while inland populations were uneven-aged. Recruitment appears to follow periods of disturbance; infrequent fire in the inland populations and continuous sand movement on the coast are factors initiating recruitment.     

Complex patterns of food allocation in asynchronously hatching broods of crimson rosellas

In birds, asynchronous hatching typically leads to lower growth and survival of last-hatched chicks. However, all crimson rosella Platycercus elegans, chicks grow at the same rate, although first-hatched chicks can be as much as seven times heavier than last-hatched chicks at the end of hatching. We examined the delivery and distribution of food to 18 rosella broods by videotaping feeds and simultaneously recording mass changes in the nestbox using a digital balance. Parents visited the nest infrequently and delivered loads of up to 25% of their body weight during a feeding visit. Male rosellas consistently delivered larger loads and consequently had higher feeding rates (g/h) than females. Parents distributed food between chicks by direct regurgitation in a series of up to 51 food transfers. Overall, chicks of all hatching ranks received equal numbers of transfers, but parents differed in how they distributed food within the brood. Males fed first-hatched chicks more than last-hatched chicks, whereas females distributed food equally to all chicks. Selective feeding of small chicks might be costly to females since they delivered food more slowly than males and spent more time in the nestbox. Thus female rosellas may invest more in current reproduction than males. Parents also distributed food differently to male and female chicks. Large males were fed more than all other nestlings, while female nestlings were fed equally irrespective of size. This study confirms that complex patterns of parental allocation occur in wild populations. Copyright 1999 The Association for the Study of Animal Behaviour.     

Male chicks are more costly to rear than females in a monogamous seabird, the Common Murre

Feeding rates and mass loss during chick rearing were comparedfor individually marked parents of male and female Common Murre(Uria aalge) chicks at Great Island, Newfoundland, Canada, from1997–2001. Both parents in this socially monogamous seabirdspecies share parental care duties until colony departure, afterwhich the single chick is fed only by its father. Because murresprovision their single chicks with one clearly visible fishper trip, it is possible to accurately determine whether parentsdifferentially feed male and female chicks. Based on slightlygreater mass of males in adulthood, possibly favored by sexdifferences in breeding roles, we predicted that male nestlingswould be fed more than females. Fathers' feeding rate to sons,but not daughters, increased with chick age, whereas maternalfeeding rate increased with chick age for both sexes. When year-correctedfeeding rates of pairs rearing both sexes were compared, bothmothers and fathers fed their sons significantly more than theirdaughters in the later part of the chick-rearing period. Moreover,parents rearing male chicks lost mass at a significantly higherrate than those rearing females. There was no difference infledging age for sons and daughters. These results indicatethat differential parental allocation occurs and has measurablecosts even in a species with only slight adult sexual dimorphism.     

The use of supplementary foods by Australian magpies Gymnorhina tibicen: Implications for wildlife feeding in suburban environments

Abstract The provision of foods to wild birds is an extremely common practice among householders throughout the Western world. Nonetheless, concerns over potential impacts of the practice are currently being raised, including the possibility that some species may become reliant on human‐provided food. We compared the foraging and breeding ecology of pairs of fed and unfed Australian magpies Gymnorhina tibicen living in suburban environments in Brisbane, Queensland, Australia. Detailed behavioural observations of both foraging and chick provisioning were made for males and females of both groups throughout the breeding season. Natural foods dominated the diets of both fed and unfed magpies, making up 76% and 92% of all items consumed respectively. During the morning, fed magpies obtained fewer food items during ground foraging than did unfed birds, apparently because they visited suburban feeding stations more often during the early part of the day. At other times, the amount of food items obtained during foraging was similar for both fed and unfed birds. Magpies utilizing suburban feeding stations started all breeding activities significantly earlier than unfed magpies, except during the fledgling phase. Both fed and unfed magpie parents provisioned their chicks predominantly with natural food. Magpies were not reliant or dependent on supplementary food provided by wildlife feeders at any time during the breeding season. Although many magpies did utilize suburban feeding stations extensively, they continued to forage for and provision their chicks with natural food.     

Sex‐specific mass loss in chick‐rearing South Polar Skuas Stercorarius maccormicki– stress induced or adaptive?

Mass loss of chick‐rearing birds can be the direct consequence of physiological stress (reproductive stress hypothesis) or an adaptive mass adjustment in response to the increased demands on flight efficiency during the flight‐intensive chick‐rearing period (adaptive mass loss hypothesis). To test which of these hypotheses best explains mass loss in South Polar Skuas Stercorarius maccormicki rearing chicks, a food supplementation experiment was carried out in the austral summer 2000/01 at King George Island, Antarctica. Half of the breeding pairs were fed about 20% of the chick's daily energy demand every second day and chick growth and adult nest attendance were recorded. Parents were caught at the start and the end of chick‐rearing to calculate adult mass loss. Male parents of food‐supplemented pairs attended their nest territories more than control males but females kept their attendance constant. Chick growth was only minimally affected and the treatment probably had no fitness consequences. Male Skuas in control pairs had a higher deviation from the body size–mass regression at the end of chick‐rearing compared with the start, supporting the stress hypothesis, whereas female deviation remained unchanged. Males of food‐supplemented pairs were heavier than unsupplemented males at the end of the breeding cycle but not significantly so. Food‐supplemented females were lighter at the end, supporting the adaptive mass loss hypothesis. Adult mass loss is thus best explained by the reproductive stress hypothesis in males but by the adaptive mass loss hypothesis in females. However, the two hypotheses are not mutually exclusive and the results do not exclude the possibility that mass loss in females is stress‐induced but the amount of mass lost is an adaptive adjustment to the reliability of the food supply. The finding that members of a breeding pair may follow different strategies of mass adjustment has implications for the use of mass loss as an index of parental effort. Without knowing which strategy each sex has adopted it is of little use to compare mass loss between parents.     

Sex-biased aggression and male-only care at sea in Brünnich's Guillemots Uria lomvia and Razorbills Alca torda

In several groups in the order Charadriformes, biparental care is followed by a period of male‐only care. Several hypotheses attempting to explain extended male parental care in shorebirds do not fit the Alcini. In a previous study of Brünnich’s Guillemots Uria lomvia and Razorbills Alca torda, we did not find support for female‐biased parental effort at the breeding site that would lead to males being in better condition to care for chicks at sea. However, in both species, males spent more off‐duty time at the breeding site than females, suggesting greater involvement in the defence of egg or chick, breeding site and mate. We predicted that there would be a male bias in size and aggressive behaviour associated with parental roles. To test this, body size and aggression of attending male and female Brünnich’s Guillemots and Razorbills were measured during incubation and brooding on the Gannet Islands, Labrador. Parental aggression was measured using natural observations of all agonistic interactions and, in Razorbills only, in situ responses to presentations of a predator model. In both species, males were significantly larger than females in culmen and gape length. Guillemot males initiated agonistic interactions more frequently than females during incubation. In contrast, female Guillemots were subjected to aggression more frequently than males and as a result were involved in more fights. In addition, the few chicks that were seen to die were being attended by single females. During the brooding period, Razorbill males responded aggressively to intruders more frequently than females, made more aggressive responses than females, and responded aggressively more frequently and more intensely than females to a predator model. In both species there was a similar male bias in morphology and behaviour that is consistent with male parents being more capable of protecting their chick, a probable advantage to chick survival during the uniparental care phase of some Charadriformes.     

Correlations between body size, defensive behaviour and reproductive success in male Little Blue Penguins Eudyptula minor: implications for female choice

Field studies exploring relationships among body size, defensive behaviour (including vocalizations) and reproductive success in male Little Blue Penguins Eudyptula minor were conducted to identify behavioural and physical attributes that might provide females or opponents with information on male quality. Head length provided a simple estimator of body size in both sexes. Nests defended by large males produced eggs and chicks earlier in the breeding season, and contained chicks that grew more quickly than chicks in nests defended by small males. In contrast, female body size did not affect any of the reproductive parameters we measured. The pitch of male defensive calls varied significantly with male size, so females (or opponents) could potentially gauge male size by assessing the calls. Large males were also more likely to respond vocally to simulated nest intrusions. As a result, by 'keeping company' with prospective partners early in the breeding season, females can obtain information about males that would allow them to select better partners and improve their own reproductive potential.     

Sex differences in responsiveness to begging in a cooperative mammal

In species where young are provisioned by both parents, males commonly contribute less to parental care than females, and are less responsive to variation in begging rates. Similar differences in the care of young occur among adults in cooperative breeders, but fewer studies have investigated whether these are associated with differences in responsiveness. Here, we present results from a playback experiment investigating responsiveness to begging in the meerkat (Suricata suricatta), a cooperatively breeding mammal. Although increased begging rate raised the feeding rate of adults of both sexes, there was no consistent tendency for females to be more responsive than males. However, when we examined changes in the proportion of food items found that were fed to pups (generosity), we found that females were more responsive than males to increased begging rate. These results can be explained in terms of sex differences in dispersal: in meerkats, females are philopatric and receive considerable benefits from investing in young, both directly, by increasing group size, and indirectly, by recruiting helpers if they inherit the breeding position. In addition, they emphasize that generosity provides a more sensitive measure of responsiveness to begging than feeding rate, as it accounts for variation in foraging success.     

Male‐skewed adult sex ratio,survival, mating opportunity and annual productivity in the Snowy Plover Charadrius alexandrinus

Sex differences in adult mortality may be responsible for male‐skewed adult sex ratios and male‐skewed parental care in some birds. Because a surplus of breeding males has been reported in serially polyandrous populations of Snowy Plover Charadrius alexandrinus, we examined sex ratio, early‐season nesting opportunities, adult survival and annual reproductive success of a Snowy Plover population at Monterey Bay, California. We tested the hypotheses that male adult survival was greater than female survival and that a sex difference in adult survival led to a skewed adult sex ratio, different mating opportunities and different annual productivity between the sexes. Virtually all females left chicks from their first broods to the care of the male and re‐nested with a new mate. As a result, females had time to parent three successful nesting attempts during the lengthy breeding season, whereas males had time for only two successful attempts. Among years, the median population of nesting Plovers was 96 males and 84 females (median difference = 9), resulting in one extra male per eight pairs. The number of potential breeders without mates during the early nesting period each year was higher in males than in females. Adult male survival (0.734 ± 0.028 se) was higher than female survival (0.693 ± 0.030 se) in top‐ranked models. Annually, females parented more successful clutches and fledged more chicks than their first mates of the season. Our results suggest that in C. alexandrinus a sex difference in adult survival results in a male‐skewed sex ratio, which creates more nesting opportunities and greater annual productivity for females than for males.     

Causes of reduced clutch size in a tidal marsh endemic

We tested three hypotheses of clutch size variation in two subspecies of the swamp sparrow (Melospiza georgiana georgiana and M. g. nigrescens). Swamp sparrows follow the pattern of other estuarine endemics, where clutch size is smaller among tidal salt marsh populations (M. g. nigrescens) than their closest inland relatives (M. g. georgiana). Our results support predation risk and temperature, but not adult survival, as explanations of this pattern in swamp sparrows. Coastal nests were twice as likely to fail as inland nests, and parental activity around the nest site was positively related to clutch size at both sites. When brood size was controlled for, coastal adults visited nests less often and females vocalized less frequently during visits than inland birds, which may decrease nest detectability to predators. Coastal parents waited longer than inland birds to feed offspring in the presence of a model nest predator, but there was no difference in their response to models of predators of adults, as would be expected if coastal birds possessed increased longevity. Additionally, coastal females laid more eggs than inland females over a single season, following a within-season bet-hedging strategy rather than reducing within-season investment. Coastal territories experienced ambient air temperatures above the physiological zero of egg development more often, and higher temperatures during laying correlated with smaller clutches and increased egg inviability among coastal birds. Similar effects were not seen among inland nests, where laying temperatures were generally below physiological zero. Both subspecies showed an increase in hatching asynchrony and a decrease in apparent incubation length under high temperatures. Coastal individuals, however, showed less hatching asynchrony overall despite higher temperatures. Both air temperatures during laying and predation risk could potentially explain reduced clutch size in not only coastal plain swamp sparrows, but also other tidal marsh endemics.     

Ontogeny of chick behavior: a tool for monitoring the growth and development of lesser adjutant storks

We studied the behavioral development of seven lesser adjutant stork (Leptoptilos javanicus) chicks from hatching to fledging over three breeding seasons at the Bronx Zoo. We developed an ethogram and compared the rate at which behaviors appeared in relation to brood size, sex, and the conditions in which the chicks were raised by their parents. Although sample sizes were small, there seem to be sex‐related differences in the rate at which behaviors develop, with females developing more rapidly than males. Larger clutch size may be associated with slower growth rate because the single male developed faster than the two males in 2004. The slowest growth rate, observed in a single male chick in 1999, was most likely owing to nutritional deficiencies and other health complications. More research is needed, but these results can be used to help evaluate the age and health of lesser adjutant stork chicks in captivity and in the wild. Zoo Biol 26:533–538, 2007. © 2007 Wiley‐Liss, Inc.     

Effects of recruiting age on senescence, lifespan and lifetime reproductive success in a long-lived seabird

Theories of ageing predict that early reproduction should be associated with accelerated reproductive senescence and reduced longevity. Here, the influence of age of first reproduction on reproductive senescence and lifespan, and consequences for lifetime reproductive success (LRS), were examined using longitudinal reproductive records of male and female blue-footed boobies (Sula nebouxii) from two cohorts (1989 and 1991). The two sexes showed different relationships between age of first reproduction and rate of senescent decline: the earlier males recruited, the faster they experienced senescence in brood size and breeding success, whereas in females, recruiting age was unrelated to age-specific patterns of reproductive performance. Effects of recruiting age on lifespan, number of reproductive events and LRS were cohort- and/or sex-specific. Late-recruiting males of the 1989 cohort lived longer but performed as well over the lifetime as early recruits, suggesting the existence of a trade-off between early recruitment and long lifespan. In males of the 1991 cohort and females of both cohorts, recruiting age was apparently unrelated to lifespan, but early recruits reproduced more frequently and fledged more chicks over their lifetime than late recruits. Male boobies may be more likely than females to incur long-term costs of early reproduction, such as early reproductive senescence and diminished lifespan, because they probably invest more heavily than females. In the 1991 cohort, which faced the severe environmental challenge of an El Ni?o event in the first year of life, life-history trade-offs of males may have been masked by effects of individual quality.     

Seasonal sex–specific energy expenditure in breeding and non-breeding Palestine sunbirds Nectarinia osea

The timing of the chick‐rearing phase is known to have a profound effect on the reproductive success of birds. However, little is known about the energetic costs faced by the parents during different periods of the breeding season. These costs may have vital consequences for both their survival and future reproduction. In most studies, daily energy expenditure (DEE) of breeding and non‐breeding birds has been compared, without controlling for the effect of season. In the present study, we examined the energy demands of breeding compared to non‐breeding Palestine sunbirds Nectarinia osea and whether there were sex‐specific differences in DEE within and between different seasons. We predicted that DEE would be elevated when birds rear chicks, especially at cooler ambient temperatures. Time‐energy budgets were constructed for pairs of sunbirds, rearing chicks, or not breeding, in spring and summer. There were significant seasonal differences in estimates of DEE in non‐breeders that were 21% higher in spring than in summer. We attributed these to increases in non‐flight metabolic rate rather than changes in time spent on different activities. Our estimates of DEE for the birds that were rearing chicks were higher than non‐breeding adults. In females the increase in DEE when breeding, compared to when not breeding, was similar in both spring and summer, while males increased their DEE much less when breeding in spring. The differences in estimated DEE, however, were not significant between male and female birds in any season. Between seasons, female breeders had 17.1% higher DEE in spring than in summer, while male breeders showed no difference in DEE when rearing chicks in different seasons. Accordingly, our initial prediction was supported, as DEE in chick‐rearing adults was higher than in non‐breeding adults. In addition, although temperatures are lower in spring, breeding in the spring is only more costly than breeding in summer for females. Apparently, males are more flexible in reallocating their time and energy spent on different activities.     

Interactions between adult robins and chicks belonging to other pairs

Adult robins are able to discriminate between chicks hatched in their own and in other territories. The latter are much less likely to be fed and are more likely to be attacked than the adult's own young. As chicks grow older they become less likely to beg at adults other than their parents. Adults rarely feed chicks from other territories unless they have fledged young of their own. Possible causes of these different responses are discussed and it is emphasized that these need not involve individual recognition. Males appear to be more likely than females to feed chicks hatching in other territories. This could be the result of the females learning about their chicks while brooding them. Males may benefit more, on average, from feeding other pairs' chicks than do females. There has been a previous report that robins are unable to make the discriminations reported in this paper and possible reasons for this discrepancy are discussed. It is conceivable that it reflects a difference between populations.     

Reproductive biology and ecology of white‐winged trumpeters (Psophia leucoptera) and recommendations for the breeding of captive trumpeters

The reproductive biology and ecology of a wild population of white‐winged trumpeters (Psophia leucoptera) were studied in southeastern Peru from 1983 to 1987. Because little information is available about any of the trumpeter species and because trumpeters have proven difficult to breed in captivity, information relevant to breeding and management of captive trumpeters is reported in this paper. White‐winged trumpeters lived in territorial social groups that ranged in size from four to 13 individuals. A typical territorial group contained three adult males, two adult females, and several sexually immature offspring, but smaller temporary groups sometimes formed for the duration of the breeding season. Only the dominant female contributed eggs to the clutch, and all adult males in the group competed to obtain copulations with her. Eggs were laid in elevated nesting cavities and no nest was constructed. The average clutch size was three eggs and incubation was not begun until the final egg was laid. The dominant male and female shared most of the incubation duties, but subordinate males covered approximately 15% of the incubation shifts. Eggs hatched approximately 27 days after incubation was begun and chicks left the nesting cavity the day after they hatched. Chicks were completely dependent on older birds to feed them for their first 3 weeks and then gradually began to feed themselves more and more food. The subordinate adult males fed chicks the most food, the dominant male and female and older offspring fed chicks an intermediate amount, and the subordinate adult female fed chicks the least. Young chicks behaved aggressively toward each other but were separated by adults before they injured each other. If at least one chick from the clutch survived, trumpeters did not breed again until the beginning of the next breeding season the following year. Zoo Biol 19:65–84, 2000. © 2000 Wiley‐Liss, Inc.     

Parental care of the Common Tern Sterna hirundo

Parental care activities of male and female Common Terns Sterna hirundo were recorded over two breeding seasons. Males and females exhibited distinct parental roles throughout a breeding bout. Courtship feeding by males was extensive prior to and during egg-laying, but declined with the onset of incubation. Females performed significantly more incubation behaviour than males although both sexes spent equal time attending at the nest site. During the chick stage, females spent significantly more time on the territory than did males. Chick feeding was largely the responsibility of the male; males fed chicks at a rate approximately three times higher than that of females. In addition, whereas females showed no trend in the size of fish delivered to chicks relative to chick age, the size of fish delivered by males increased with chick age. Courtship feeding activities and extensive chick feeding contributions by male Common Terns appear to outweigh parental contributions by females, contrary to predictions for a monogamous species.     

Reversal of height dimorphism promotes pollen and seed dispersal in a wind-pollinated dioecious plant

Variation in the timing of reproductive functions in dioecious organisms may result in adaptive changes in the direction of sexual dimorphism during the breeding season. For plants in which both pollen and seeds are wind-dispersed, it may be advantageous for male plants to be taller when pollen is dispersed and female plants to be taller when seeds are dispersed. We examined the dynamics of height dimorphism in Rumex hastatulus, an annual, wind-pollinated, dioecious plant from the southern USA. A field survey of seven populations indicated that females were significantly taller than males at seed maturity. However, a glasshouse experiment revealed a more complex pattern of height growth during the life cycle. No dimorphism was evident prior to reproduction for six of seven populations, but at flowering, males were significantly taller than females in all populations. This pattern was reversed at reproductive maturity, consistent with field observations. Males flowered later than females and the degree of height dimorphism was greater in populations with a later onset of male flowering. We discuss the potential adaptive significance of temporal changes in height dimorphism for pollen and seed dispersal, and how this may be optimized for the contrasting reproductive functions of the sexes.     

The effect of parental age,experience and historical reproductive success on wandering albatross (Diomedea exulans) chick growth and survival

Growth and survival of altricial young are influenced by their parents’ abilities to invest in a breeding attempt. As a result, chick growth and survival in one breeding season may be indicative of their parents’ long-term reproductive potential. To determine whether variation in long-term reproductive success is driven by differential breeding investment, parental care and chick growth in wandering albatrosses (Diomedea exulans) were correlated with parental historical reproductive success. Effects of age and breeding experience (determined from past breeding attempts) and pre-laying body condition (mass–size indices) on chick growth and survival also were tested. Longer brooding of chicks increased their survival, but length of chick brooding did not differ between historically unproductive and successful breeders. Past reproductive success also was not correlated with chick growth rates or fledging mass or size. Chick brooding period, chick growth rates, final mass and size were independent of parental body condition. Older and more experienced parents brooded chicks for longer and their chicks grew faster, supporting previous findings that breeding competence is a learnt skill. Chick care and growth characteristics differed more between than within pairs, suggesting that differences in these characteristics are driven by variation among pairs.     

Prolactin and parental behavior in Adélie penguins: effects of absence from nest, incubation length, and nest failure

Adélie penguin (Pygoscelis adeliae) males and females, nesting in Antarctica, alternate attendance at the nest with absences of many days to forage at sea. We investigated the importance of tactile input from egg and chicks on prolactin levels by observing nest attendance patterns and obtaining blood samples (1) during the first nest exchange of the incubation stage, (2) from birds whose incubation period was artificially increased or decreased by about 10 days, and (3) from birds whose nests had failed. Prolactin levels in females after 8 to 11 days of absence from the breeding colony did not differ from those in incubating males and did not change after females resumed incubation. Moving eggs between nests resulted in nests in which chicks hatched after about 26, 36 (normal), or 46 days. Duration of incubation did not affect prolactin levels in the parents measured during incubation, at the pip stage, hatch stage, or early brood stage. Adults first left their chicks unguarded on about the same calendar date, regardless of chick age. However, chicks from long incubation nests averaged 8 days younger when they were left unguarded than chicks from control or short-incubation nests. In females, there was no effect of nest failure on prolactin levels. In males, prolactin levels were slightly lower after nest failure than in males tending nests. Testosterone was significantly higher in males after nest failure than in males still tending nests. Prolactin is elevated in Adélie penguins as part of the program of cyclical hormonal changes that accompany the lengthy reproductive season and is relatively independent of tactile input. Sustained prolactin secretion is probably required for the maintenance of parental behavior in offshore feeding species that must be absent from the nest for many days at a time.