Annual variation in reproductive parameters of Adélie penguins at Edmonson Point,Victoria Land,Antarctica

Annual egg and chick production and breeding success at the Adélie penguin (Pygoscelis adeliae) colony Edmonson Point (74°21′S–165°10′E), Victoria Land, is presented for eight breeding seasons between 1995 and 2005. During this period the colony consisted of 10–13 subcolonies and averaged 2098 ± 278 breeding pairs. A sample of over 100 nests (114–150), belonging to two subcolonies, was monitored each year. Some breeding parameters remained constant while others showed substantial annual variation. Laying date showed little variation, and laying was highly synchronous: 82.5% of clutches were initiated in a 10-day period, 9–18 November. In contrast, clutch size (1.77–1.97) and incubation period (34.4 ± 2.5) varied significantly. Variation among years was also recorded in hatching success (from 58 to 86%) and breeding success. This last parameter, measured as number of chicks reared to crèche per nest with eggs, varied between 0.34 and 0.97.     

Population variability in heat shock proteins among three Antarctic penguin species

Heat shock proteins (HSPs) are synthesised under stressful conditions such as exposure to elevated temperatures, contamination, free radicals, UV light or pathophysiological states resulting from parasites and/or pathogens. HSPs function to protect cells by means of modulation of protein folding. In Antarctica, these proteins have been studied in such organisms as protozoa and fishes, without attention to geographical variation. We studied the variation of HSP70 and HSP60 levels in Gentoo, Adelie and Chinstrap penguins among different populations along the Antarctic Peninsula from King George Island (62°15′S) to Avian Island (67°46′S). Our results show that the northern population of Gentoo penguin showed higher levels of HSP70 and HSP60 than the southern population. High temperature, human impact and immunity as a proxy for parasites and diseases in northern locations could explain such variation. Adelie penguin only showed significant geographical variation in HSP70, increasing north to south, a pattern perhaps related to increased UV radiation and decreased temperatures from north to south. Chinstrap penguin shows no population differences in the variation in neither HSP70 nor HSP60, although HSP70 showed marginally significant differences. Sexual differences in the level of these proteins are also discussed.     

Relationship between reversed sexual dimorphism,breeding investment and foraging ecology in a pelagic seabird,the masked booby

Reversed sexual dimorphism (RSD) may be related to different roles in breeding investment and/or foraging, but little information is available on foraging ecology. We studied the foraging behaviour and parental investment by male and female masked boobies, a species with RSD, by combining studies of foraging ecology using miniaturised activity and GPS data loggers of nest attendance, with an experimental study where flight costs were increased. Males attended the chick more often than females, but females provided more food to the chick than males. Males and females foraged during similar periods of the day, had similar prey types and sizes, diving depths, durations of foraging trips, foraging zones and ranges. Females spent a smaller proportion of the foraging trip sitting on the water and had higher diving rate than males, suggesting higher foraging effort by females. In females, trip duration correlated with mass at departure, suggesting a flexible investment through control by body mass. The experimental study showed that handicapped females and female partners of handicapped males lost mass compared to control birds, whereas there was no difference for males. These results indicate that the larger female is the main provisioner of the chick in the pair, and regulates breeding effort in relation to its own body mass, whereas males have a fixed investment. The different breeding investment between the sexes is associated with contrasting foraging strategies, but no clear niche differentiation was observed. The larger size of the females may be advantageous for provisioning the chick with large quantities of energy and for flexible breeding effort, while the smaller male invests in territory defence and nest guarding, a crucial task when breeding at high densities. In masked boobies, division of labour appears to be maximal during chick rearing—the most energy-demanding period—and may be related to evolution of RSD.     

PARENTAL QUALITY AND SELECTION ON EGG SIZE IN THE MAGELLANIC PENGUIN

We examined the relative contributions of egg size and parental quality to hatching success, fledging success, and chick growth in the Magellanic penguin (Spheniscus magellanicus) be exchanging clutches between nests to reduce the covariation between egg and parental factors. Among control nests, fledging success increased slightly with egg size. However, the effect of egg size independently of parental quality was limited to an influence on chick mass and size for the first 10 days post-hatching. In contrast, attributes of the parents influenced nesting success and chick size at fledging, independently of the egg size actually raised. We suggest that the common occurrence of a positive phenotypic correlation between egg size and fledging success is due to two factors: (1) adults laying large eggs tend to be of higher quality; and (2) to the extent that egg size does influence early survival independently of parental quality, the effect on survival is due to a maternal effect on egg composition rather than an inherent effect of egg size.     

Estimating the relative abundance of emperor penguins at inaccessible colonies using satellite imagery

Emperor penguin (Aptenodytes forsteri) populations are useful environmental indicators due to the bird’s extreme reliance on sea ice. We used remote sensing technology to estimate relative adult bird abundance at two inaccessible emperor penguin colonies in the Ross Sea, Antarctica. We performed supervised classification of 12 panchromatic satellite images of the seven known Ross Sea colonies. We used regression to predict adult bird counts at the inaccessible colonies by relating the number of pixels classified as “penguin” in the satellite images of the accessible colonies to corresponding known adult bird counts from aerial photographs or ground counts. While our analysis was hampered by excessive guano and shadows, we used satellite imagery to differentiate between relatively small (<3,000 adult birds) and larger colonies (>5,000 adult birds). Remote sensing technology is logistically less intense and less costly than aerial or ground censuses when the objective is to document penguin presence and/or large emperor penguin population changes (e.g., catastrophic changes). Improvements expected soon in the resolution of the satellite images should allow for more accurate abundance estimates.     

Reproductive investment and parental roles in Sabine's gulls<Emphasis Type="Italic"> Xema sabini</Emphasis>

More than 90% of avian species exhibit biparental care, though parental activities are often shared unequally between the members of a pair. Among gull species (Laridae), males and females generally share parental activities, although there appear to be considerable differences between species in the relative contribution of each sex. This study examined the behaviour of male and female Sabine's gulls (Xema sabini) during the incubation period and immediately post-hatch; particularly the amount of time they each invested in breeding activities. Although considered an aberrant species in aspects of behaviour and biology, the Sabine's gull showed a high reproductive investment by both sexes, as other gull species do. Males fed females prior to egg laying and contributed equally to incubation and chick provisioning, and females contributed equally to nest defence. Overall, there was no difference between the sexes in the extent of their contributions to parental care, although there was considerable individual variation within pairs. Sabine's gulls are Arctic breeders and the extent of their contributions to parental activities could have been influenced by their extreme breeding environment and short breeding season.     

The variation in reliability of individual vocal signature throughout ontogenesis in the red-crowned crane Grus japonensis

Distinctiveness of chicks’ calls may arise in ontogenesis when parents can confuse their own and alien chicks, leaving their nests and forming crèches or flocks. It is unknown, however, whether the individual vocal signature retains further in ontogenesis or relaxes when the necessity in the parental care disappears. In this paper, we study the inter- and intra-individual variations of the acoustic parameters in chicks’ calls in the red-crowned crane Grus japonensis, the species with prolonged development enveloping three stages: territorial under parental care, in flocks under parental care and in flocks self-independently. We found, that discriminability of chicks’ calls increased significantly to the second stage, characterized by the maximum risk for parents to confuse the own and alien chicks, and significantly decreased to the third stage, when the needs in the parent–chick vocal recognition disappeared. Our data agree with a hypothesis that the individual distinctiveness decreases in the absence of necessity in accuracy of parent–chick recognition.     

Analysis of the sequence variations in the Mhc DRB1-like gene of the endangered Humboldt penguin (Spheniscus humboldti)

The Major Histocompatibility Complex (Mhc) genomic region of many vertebrates is known to contain at least one highly polymorphic class II gene that is homologous in sequence to one or other of the human Mhc DRB1 class II genes. The diversity of the avian Mhc class II gene sequences have been extensively studied in chickens, quails, and some songbirds, but have been largely ignored in the oceanic birds, including the flightless penguins. We have previously reported that several penguin species have a high degree of polymorphism on exon 2 of the Mhc class II DRB1-like gene. In this study, we present for the first time the complete nucleotide sequences of exon 2, intron 2, and exon 3 of the DRB1-like gene of 20 Humboldt penguins, a species that is presently vulnerable to the dangers of extinction. The Humboldt DRB1-like nucleotide and amino acid sequences reveal at least eight unique alleles. Phylogenetic analysis of all the available avian DRB-like sequences showed that, of five penguin species and nine other bird species, the sequences of the Humboldt penguins grouped most closely to the Little penguin and the mallard, respectively. The present analysis confirms that the sequence variations of the Mhc class II gene, DRB1, are useful for discriminating among individuals within the same penguin population as well those within different penguin population groups and species.The nucleotide sequence and amino acid sequence data reported in this paper have been submitted to the DDBJ database and have been assigned the accession numbers AB088371–AB088374, AB089199, AB154393–AB154399, and AB162144.     

Foraging areas of Wilson’s storm-petrel Oceanites oceanicus in the breeding and inter-breeding period determined by stable isotope analysis

To understand the year-round ecology of seabirds it is necessary not only to study the birds in their breeding grounds, but also to gain information about their movements during the inter-breeding period. Especially for the smaller procellariiform species, such studies are still scarce, mainly due to methodological problems. The recovery rates of banded birds are low and satellite tracking devices still far too heavy to equip these small birds. Here, we present data on foraging areas of Wilson’s storm-petrel Oceanites oceanicus inferred from stable isotope analysis. We compared ratios of δ¹³C and δ¹⁵N between different life-history stages and between the breeding and inter-breeding period. Samples of adult and chick feathers, chick down and egg-white were taken between 1996 and 2005 on King-George-Island, South Shetland Islands. δ¹³C values can be clearly distinguished between the breeding and inter-breeding period. During the inter-breeding period, most pre-breeders foraged in the same area as breeders, but four pre-breeders were found to forage in latitudes north of the Subtropical Front. In the 2002 inter-breeding period adult birds wintered further north than in 2003, which is in line with the different locations of food rich frontal systems in these years. We show that isotope ratios of both δ¹³C and δ¹⁵N increase from egg white, over chick down to chick feathers. We suggest that this isotopic change, due to a change in both foraging location and diet between egg production and chick feeding, may be used to trace the shift from the use of maternal resources from the egg to the uptake of nutrients from the diet.     

Cycles of <Emphasis Type="Italic">Euphausia superba</Emphasis> recruitment evident in the diet of Pygoscelid penguins and net trawls in the South Shetland Islands,Antarctica

Size and sex of Antarctic krill taken from chinstrap and gentoo penguin diet were compared to those from scientific net surveys in the South Shetland Islands from 1998 to 2006 in order to evaluate penguin diet as a sampling mechanism and to look at trends in krill populations. Both penguin diet and net samples revealed a 4–5 year cycle in krill recruitment with one or two strong cohorts sustaining the population during each cycle. Penguin diet samples contained adult krill of similar lengths to those caught in nets; however, penguins rarely took juvenile krill. Penguin diet samples contained proportionately more females when the krill population was dominated by large adults at the end of the cycles; net samples showed greater proportions of males in these years. These patterns are comparable to those reported elsewhere in the region and are likely driven by the availability of different sizes and sexes of krill in relation to the colony.     

Breeding site fidelity in penduline tit Remiz pendulinus in Southern Hungary

Birds move between breeding locations to gain a better territory, avoid competition or reduce the deleterious effect of inbreeding. We investigated breeding site fidelity in a small European passerine, the penduline tit (Remiz pendulinus). This species has an exceptionally diverse breeding system, in which both males and females may have up to 5–7 mates in a single breeding season, and the eggs are incubated by a single parent: either the male or the female. We investigated the movements of males and females within three breeding seasons in Southern Hungary (2002–2004). Males moved for shorter distances between breeding sites (116 m, 63–333 m; median, lower quartile–upper quartile) than females (942 m, 415–2,382 m). Movements of males and females were consistent between years, and they were repeatable between subsequent nests of males, but not of females. Taken together, our results suggest that adult male penduline tits are more site-faithful than adult females. We suggest that this difference has an implication on their breeding ecology since male parental behaviour (desert/care) is expected to be influenced by local mating opportunities, whilst female parental behaviour is likely to depend on the mating opportunities in a large area around their breeding site.     

Low MHC variation in the endangered Galápagos penguin (<Emphasis Type="Italic">Spheniscus mendiculus</Emphasis>)

The major histocompatibility complex (MHC) is one of the most polymorphic regions of the genome, likely due to balancing selection acting to maintain alleles over time. Lack of MHC variability has been attributed to factors such as genetic drift in small populations and relaxed selection pressure. The Galápagos penguin (Spheniscus mendiculus), endemic to the Galápagos Islands, is the only penguin that occurs on the equator. It relies upon cold, nutrient-rich upwellings and experiences severe population declines when ocean temperatures rise during El Niño events. These bottlenecks, occurring in an already small population, have likely resulted in reduced genetic diversity in this species. In this study, we used MHC class II exon 2 sequence data from a DRB1-like gene to characterize the amount of genetic variation at the MHC in 30 Galápagos penguins, as well as one Magellanic penguin (S. magellanicus) and two king penguins (Aptenodytes patagonicus), and compared it to that in five other penguin species for which published data exist. We found that the Galápagos penguin had the lowest MHC diversity (as measured by number of polymorphic sites and average divergence among alleles) of the eight penguin species studied. A phylogenetic analysis showed that Galápagos penguin MHC sequences are most closely related to Humboldt penguin (Spheniscus humboldti) sequences, its putative sister species based on other loci. An excess of non-synonymous mutations and a pattern of trans-specific evolution in the neighbor-joining tree suggest that selection is acting on the penguin MHC.     

An observation of between-mates feeding behaviour in chick-guarding chinstrap penguins

Courtship feeding has not been reported for chinstrap penguins (Pygoscelis antarctica). However, we observed a male chinstrap penguin fed to his mate during the guard stage of chick. After being fed, the female regurgitated the food to her chick. Our observation suggests that the chicks’ behaviour of begging for food may be retained in adults, but it is usually restrained.     

Consequences of chronic infections with three different avian malaria lineages on reproductive performance of Lesser Kestrels (Falco naumanni)

We studied the consequences of chronic infections by three different lineages of avian malaria, two Plasmodium (RTSR1, LK6) and one Haemoproteus (LK2), on reproductive performance of Lesser Kestrels (Falco naumanni). Malaria infections in male and female parents had no effect on clutch size, hatching success or nesting success. However, when only successful nests were considered, we found that males parasitized by LK6 raised a lower number of fledglings, suggesting that the level of parental effort by males may be limited by this particular lineage of Plasmodium. This effect was not evident in females, probably due to the higher investment of males during the chick rearing period in this species. Overall, we have found that chronic stages of specific malaria lineages have certain negative consequences on host reproductive performance, highlighting the importance of considering genetic differences among malaria parasites to study their consequences on natural bird populations.     

Ranging Behavior and Possible Correlates of Pair-Living in Southeastern Avahis (Madagascar)

Researchers have proposed several hypotheses to explain pair-living in primates. In particular, when males are not involved in direct parental care, pair-living may be related to female dispersal, infanticide prevention, or male mate/resource defense. We aimed to evaluate, through a better understanding of the ranging patterns of avahis, which hypotheses may best account for pair-living in these nocturnal lemurs. We collected focal observations over 26 nights, June–September 2004, in a littoral forest (Sainte Luce, southeastern Madagascar) on 4 adult radiocollared avahis [Avahi laniger (Tattersall, I. (1982). The Primates of Madagascar. Columbia University Press, New York.) or Avahi meridionalis (Zaramody in Primate Reports 74:9–22, 2006)]. We followed 2 males and 2 females from 2 groups: B, a male-female couple, and A, comprising the parental pair and 1-yr-old female offspring. The adult females birthed in August. We recorded resting and feeding tree points (via global positioning system) for home range calculation through minimum convex polygon and kernel methods. We provide the first quantitative information on the ranging behavior of 2 male and 2 female avahis. Home range/daily path length values (means) are higher than the ones previously reported for the same and other folivorous pair-living lemur species. On average, the 2 females spent more time feeding and traveled shorter distances than the 2 males did. Male–female cohesion (mean values), possibly enhanced by offspring presence, was higher in A and after births than in B and before births, respectively. Although male avahis may be forced into pair-living owing to energy constraints related to size, locomotion, and diet, females might accept pair-living in exchange for indirect territory defense and mate guarding.     

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.     

Breeding biology and provisioning of nestling snow buntings in the Canadian High Arctic

We examined basic breeding biology, as well as the effects of parental sex, brood size, nestling age, and nest habitat on foraging distances and parental food delivery rates of snow buntings (Plectrophenax nivalis) on Devon Island, NT, Canada, in the summers of 2003–2006. Clutch sizes and initiation dates were similar to those found in previous studies, although nest density was much higher. Feeding rates and foraging distances of buntings differed depending on the size and age of the brood, but were similar between nest habitats. Unlike in earlier studies, we found that male buntings made fewer feeding visits, but traveled longer distances to forage for food than females. Whether these differences between the sexes relate to reducing competition for prey, or to differing sex roles in parental care due to the poorly insulated nest cavities of this species, remains to be determined.     

The benefit to some minnows of spawning in the nests of other species

Synopsis Fishes that act as nest associates spawn simultaneously with nest-building hosts and then abandon their eggs. The proposed benefit for this behavior is increased brood survivorship, arising from the physical environment provided by the nest or the parental care provided by the host. Field and enclosure experiments indicated that associates benefit from the parental care provided by the host, and not from the physical environment provided by the nests of hosts. This information, along with the effect of nest association on host reproductive success, is necessary before the nature of this nesting symbiosis can be characterized.     

Meiotic behaviour of Un,D and R genomes in the amphiploid Aegilops ventricosa -Secale cereale and the parental species

Summary Meiotic pairing frequencies of the Un and D genomes of Ae. ventricosa and the R of S. cereale could be easily established at metaphase I in Aegilops ventricosa — Secale cereale amphiploid plants as well as in its parental species by using the C-banding technique procedure. The results show a high diminution of chromosome pairing for all genomes in the amphiploid with respect to its parental species probably due to C-heterochromatin content and/or genotypic or cryptic interactions between the three genomes.     

Parental quality and egg size affect chick survival in a precocial bird, the lapwing Vanellus vanellus

Effects of egg size and parental quality on lapwing Vanellus vanellus chick survival were studied in southwestern Sweden over 6 years. Chicks from large eggs were heavier at hatching and survived significantly better than those from small eggs. To control for the confounding effect of parental quality on egg size and chick survival, we performed a cross-fostering experiment during 2 years, exchanging clutches between nests with large and small eggs. In control clutches, chicks from large eggs survived better than those from small eggs, but we found no significant difference in chick survival between exchanged clutches. Thus, egg size did not affect chick survival independently of parental quality. Fledging success increased with parental age and/or experience, and with female body mass. Hence, both egg size and parental quality affect chick survival in the lapwing. Received: 22 February 1996 / Accepted: 30 September 1996