Genetic isolation and divergence in sexual traits: evidence for the northern rockhopper penguin Eudyptes moseleyi being a sibling species.

The taxonomic status of populations of rockhopper penguins (Eudyptes chrysocome) is still enigmatic. Northern populations differ from southern ones in breeding phenology, song characteristics and head ornaments used as mating signals. We conducted a molecular analysis using mitochondrial DNA sequencing to test if there is a gene flow barrier between northern (subtropical) populations and southern (subantarctic) populations in relation to the Subtropical Convergence, a major ecological boundary for marine organisms. Sequences of the control region and the ND2 gene were analysed in rockhopper penguins and in the macaroni penguin (Eudyptes chrysolophus), a closely related species. Genetic distances and phylogenetic analyses showed a clear split into three clades, two rockhopper clades and the macaroni penguin. Moreover, Theta(ST) and gene flow estimates also suggested genetic structuring within the northern rockhoppers. Our results add further support to the notion that the two rockhopper penguin taxa, often considered as two subspecies, can be recognized as two species E. chrysocome and E. moseleyi. The divergence in mating signals found between these two taxa seems to have occurred recently and relatively rapidly. Thus, the behavioural changes may have been enough to isolate these taxa without the need for morphological differentiation. The findings have important conservational implications, since E. moseleyi is far less abundant than E. chrysocome, but more populations may warrant an uplisting to endangered status if full species status should be recognized for more subpopulations.

Evidence that blue petrel, Halobaena caerulea, fledglings can detect and orient to dimethyl sulfide.

Procellariiform seabirds (the petrels, albatrosses and shearwaters) are recognized for their acute sense of smell. These pelagic seabirds forage over thousands of miles of ocean to find patchily distributed prey resources. Over the past decade, much headway has been made in unravelling the variety of olfactory foraging strategies that Antarctic species employ, and it is becoming clearer that olfaction plays a key role in foraging, particularly for burrow nesting species. Now we are beginning to explore how these behaviours develop in chicks. Procellariiform chicks fledge and survive the open seas without aid or instruction from a parent, but how they are able to accomplish this task is unknown. Here we explore whether chicks leave the nest pre-tuned to olfactory cues necessary for foraging. In this study, we tested the hypothesis that blue petrel chicks (Halobaena caerulea) are able to detect and orient to a foraging cue (dimethyl sulphide, DMS) used by adults without ever having experienced this odour at sea. We first established that chicks could detect DMS at a biologically relevant concentration that they will later naturally encounter at sea (<10 pmol l-1). We then performed preference tests in a Y-maze on a group of birds 1-6 days before they fledged. Sixteen out of 20 fledglings preferred DMS (e.g. DMS+propylene glycol) to a ;control' odour (propylene glycol alone). Our results suggest that chicks can detect and may already recognize DMS as an orientation cue even before they leave the nest to forage for the first time.

Orientation in the wandering albatross: interfering with magnetic perception does not affect orientation performance.

After making foraging flights of several thousands of kilometers, wandering albatrosses (Diomedea exulans) are able to pinpoint a specific remote island where their nests are located. This impressive navigation ability is highly precise but its nature is mysterious. Here we examined whether albatrosses rely on the perception of the Earth's magnetic field to accomplish this task. We disturbed the perception of the magnetic field using mobile magnets glued to the head of nine albatrosses and compared their performances with those of 11 control birds. We then used satellite telemetry to monitor their behavior. We found that the ability of birds to home specific nest sites was unimpaired by this manipulation. In particular, experimental and control birds did not show significant differences with respect to either foraging trip duration, or length, or with respect to homing straightness index. Our data suggest that wandering albatrosses do not require magnetic cues to navigate back to their nesting birds.

Penguins use the two-voice system to recognize each other.

The sound-producing structure in birds is the syrinx, which is usually a two-part organ located at the junction of the bronchi. As each branch of the syrinx produces sound independently, many birds have two acoustic sources. Thirty years ago, we had anatomical, physiological and acoustical evidence of this two-voice phenomenon but no function was known. In songbirds, often these two voices with their respective harmonics are not activated simultaneously but they are obvious in large penguins and generate a beat pattern which varies between individuals. The emperor penguin breeds during the Antarctic winter, incubating and carrying its egg on its feet. Without the topographical cue of a nest, birds identify each other only by vocal means when switching duties during incubation or chick rearing. To test whether the two-voice system contains the identity code, we played back the modified call of their mate to both adults and also the modified call of their parents to chicks. Both the adults and the chicks replied to controls (two voices) but not to modified signals (one voice being experimentally suppressed). Our experiments demonstrate that the beat generated by the interaction of these two fundamental frequencies conveys information about individual identity and also propagates well through obstacles, being robust to sound degradation through the medium of bodies in a penguin colony. The two-voice structure is also clear in the call of other birds such as the king penguin, another non-nesting species, but not in the 14 other nesting penguins. We concluded that the two-voice phenomenon functions as an individual recognition system in species using few if any landmarks to meet. In penguins, this coding process, increasing the call complexity and resisting sound degradation, has evolved in parallel with the loss of territoriality.

Reproductive endocrinology of tropical seabirds: sex-specific patterns in LH, steroids, and prolactin secretion in relation to parental care.

Plasma levels of luteinizing hormone, prolactin, testosterone, and progesterone were measured throughout breeding in masked boobies, red-footed boobies, and red-tailed tropicbirds at Europa and Tromelin Islands (Indian Ocean). LH secretion showed a dampened pattern in the three species, particularly in tropicbirds. Such specific differences may be related to the less elaborate courtship displays in tropicbirds. Testosterone levels were very low throughout breeding in all three species, particularly in boobies. Low testosterone values in boobies may be related to their year-round attendance at the colony. Prolactin secretion increased from the prelaying period until the incubation and brooding periods and declined thereafter in boobies but stayed relatively unchanged throughout the breeding cycle in tropicbirds. The relatively constant prolactin secretion in the more pelagic tropicbirds might allow them to undertake parental care despite long absences at sea. Boobies perform postfledging care with basal prolactin levels. For all species, females always have higher prolactin levels than males. This hormonal dimorphism, being more pronounced in boobies, may be associated with differences in parental care between mates.

Perceptual salience of individually distinctive features in the calls of adult king penguins.

In the king penguin, Aptenodytes patagonicus, incubation and brooding duties are undertaken alternately by both partners of a pair. Birds returning from foraging at sea find their mate in the crowded colony using acoustic signals. Acoustic recognition of the mate maintains and strengthens the mate's fidelity and favors synchronization in the different stages of reproduction. In this study it was found that the king penguin vocalizes in response to the mate's playback calls, but not to those of neighbors or unfamiliar conspecific individuals. To study individual features used by the birds for individual recognition of mates, various experimental signals consisting of synthesized modifications of the mate' s call were played back to the incubating bird. Results indicated that birds attend to the FM profile of the call, in particular its initial inflexion. The frequency modulation shape of the syllable can be assimilated to a vocal signature repeated though the different syllables of the call. King penguins pay little attention to the call' s AM envelope or its absolute frequency.

Finding a parent in a king penguin colony: the acoustic system of individual recognition.

To be fed, a king penguin, Aptenodytes patagonicus, chick must identify the call of its parents, in the continuous background noise of the colony. To study this recognition process, we played back to the chicks parental calls with acoustic parameters modified in the temporal and frequency domains. The parental call is composed of syllables (complex sounds with harmonic series) separated by pronounced amplitude declines. Our experiments with modified signals indicate that the chick's frequency analysis of the call is not tuned towards precise peak energy values, the signal being recognized even when the carrier frequency was shifted 100 Hz down or 75 Hz up. To recognize the adult, chicks used frequency rather than amplitude modulation, in particular the frequency modulation shape of the syllable. This structure is repeated through the different syllables of the call giving a distinct vocal signature. Our experiments also show that the receiver needs to perceive only a small part of the signal: the first half of the syllable (0.23 s) and the first three harmonics were sufficient to elicit recognition. The small amount of information necessary to understand the message, the high redundancy in the time and frequency domains and the almost infinite possibilities of coding provided by the frequency modulation signature permit the chick to recognize the adult, without the help of a nest site. For these reasons, the code used in the call of the king penguin can be regarded as a functional code, increasing the possibility of individual recognition in an acoustically constraining environment. Copyright 1999 The Association for the Study of Animal Behaviour.

Age-related mate choice in the wandering albatross.

We studied mate choice in the wandering albatross, Diomedea exulans, using data from 32 years of banding returns in the population of the Crozet Islands. We studied mating choices in a single year, when the Crozet Islands population was male biased (8:5, males:females). Thus, we expected that females might show great flexibility of choice of partners. Because age and experience might influence mate choice, we tested the expectation that females would choose the oldest and most experienced males for pair bonding. Pair bonds usually last until one member of the pair dies (0.3% of the birds 'divorce'), so mate choice should be especially important. We found that the ages of males and females in both displaying and bonded (breeding) pairs were significantly correlated. These age-associated pairings were not a passive phenomenon, but appeared to be due to an active process of selection of mates of similar age. First-time breeders sought mates of similar age, but preferred those with the most experience. Remating, experienced birds whose mates had died did not pair with individuals of significantly similar age, but predominantly paired with other widowed birds that, on average, were also relatively old. Mate fidelity in wandering albatrosses may be due to the cost of finding and bonding with a new mate. Pair bonds, and thus breeding, took an average of 3.2 and 2.3 years to establish, for males and females, respectively. Thus, remating exerts a potential average reproductive cost of about 15% of lifetime reproductive success. Copyright 1999 The Association for the Study of Animal Behaviour.

Endocrine correlates of parental care in an Antarctic winter breeding seabird, the emperor penguin, Aptenodytes forsteri.

Plasma levels of luteinizing hormone (LH) and prolactin associated with parental behavior were measured in the Antarctic winter breeding emperor penguin, Aptenodytes forsteri. Males exclusively incubate the egg while females exclusively brood the nonhomeothermic young; both sexes alternate in rearing the homeothermic young. Birds were sampled on arrival from the sea through egg laying, incubation, and brooding. All parent birds lost their chicks at the end of the brooding period due to harsh weather but sampling continued. In females, LH titers dropped after egg laying but levels were restored when the birds returned from the sea to brood the chicks and were not depressed by high prolactin levels. Plasma prolactin remained low in males captured on arrival and kept until the free-living males finished incubation. In breeders, prolactin secretion increased during the prelaying period when day length decreased. Prolactin levels stayed elevated in males during incubation and in brooding females returning after a 2-month absence at sea. Prolactin values were higher in brooding females than in males ending incubation or returning in late brooding. These levels did not drop after chick loss, and the sexual difference in prolactin values was maintained after breeding failure. In emperor penguins, increased prolactin secretion appears to be triggered around the time of egg laying and continues, driven by an endogenous mechanism, through incubation and brooding until rearing is completed. Prolactin secretion independent of external stimuli may have evolved in pelagic seabirds to maintain parental care despite long absences at sea from the breeding colony.

A method of independent time and frequency decomposition of bioacoustic signals: inter-individual recognition in four species of penguins.

The aim of this paper is to introduce a method for analyzing acoustic signals capable of assessing the potential for individual coding information. Signals are analysed both in the time domain (rhythm of emission of the song independent of its frequency content) and in the spectral domain (spectral content of the song independent of the rhythm of emission). The method is then applied to a comparative study of four penguin species, where the problem posed by inter-individual recognition differs from species to species. A direct relationship was shown between the potential of individual coding and the difficulty in partner identification.

Parental care and the prolactin secretion pattern in the King penguin: an endogenously timed mechanism?

The King penguin (Aptenodytes patagonicus) has been studied on Possession Island, Crozet Archipelago (46 degrees 25'S-51 degrees 45'E). It is an offshore feeder, but it breeds on land. Its breeding cycle is unusually long (about 14 months). It starts at the beginning of spring, is interrupted during 5 months of winter, and ends in the next spring. Furthermore, it is characterized by a long parental care period, of about 11 months, including the winter interruption. In fact, care given to the egg and the chick is biparental, which supposes that parental behavior includes both parents. Each parent alternates care given to the egg and to the chick on land and foraging bouts at sea. An incubation bout, or a chick care bout, is called a shift. Prolactin is the hypophyseal hormone known to be correlated with incubation and chick care. We studied the mechanism of the maintenance of prolactin during the parental care period in the King penguin, a period which is unusually long. In many species, prolactin secretion has been shown to be stimulated by the presence of eggs and/or chicks, but in the King penguin, prolactin secretion is observed throughout the entire period of parental care, despite the fact that the birds leave the egg and the chick repeatedly and for extended periods of time to feed. Prolactin levels rise significantly at the beginning of courtship; females have significantly higher prolactin levels than males during courtship, copulation, and the period of waiting for egg laying. In both sexes, prolactin levels remain high during incubation and the first part of chick rearing, before winter. Prolactin concentrations decline somewhat during the winter period of minimal parental care, but remain that level in spring when parental care starts again. The level returns to basal value during molt. Prolactin levels rise during the incubation shifts but not over the course of contact with young. Prolactin values remain high in unsuccessful breeders, possibly preventing the birds from relaying, but remain low in immature birds. These data raise questions about how prolactin secretion is controlled in this species. The hypothesis of a programmed secretion of prolactin is advanced.

Endocrine basis of the reproductive pattern of the Gentoo penguin (Pygoscelis papua): winter breeding and extended laying period in northern populations.

Changes in plasma LH, prolactin, testosterone, estradiol, and progesterone were investigated throughout moult and reproduction in free-living male and female Gentoo penguins (Pygoscelis papua) at Crozet Island (46 degrees S, 51 degrees E), where this species is able to relay after a reproductive failure. In both sexes, LH, prolactin, and steroid hormones, remained at basal levels during the moult. LH level was highest at the time of arrival at the colony for breeding and, although it decreased after courtship, it did not drop at basal value by incubation and first chick brooding period. Prolactin peaked for both chick brooding periods; replacement clutch was associated with an increased secretion of LH, whereas high prolactin levels were maintained. Testosterone, in male, and estradiol, in female, peaked during courtship I and chick brooding II; progesterone, in female, peaked during courtship I and II. These hormonal patterns are consistent with those observed in passerine species which are also able to relay after a reproductive failure. Winter breeding observed at Crozet Island might reflect the extreme adaptive capacity of Gentoo penguin species.

Plasma LH and steroid hormones in King penguin (Aptenodytes patagonicus) during the onset of the breeding cycle.

Temporal changes in plasma levels of luteinizing hormone (LH), testosterone, estradiol, and progesterone were measured throughout molt and the onset of reproduction in free-living male and female King penguins (Aptenodytes patagonicus) at Crozet Island (46 degrees S, 51 degrees E). In both sexes, LH concentrations began to rise as soon as molting ended, before the departure for refeeding at sea, suggesting that the beginning of reproductive activity is not associated with refeeding and/or courtship behavior. LH remained at high values throughout courtship and decreased to lower levels during incubation. Gonadotrophin secretion covaried with sustained plasma testosterone levels in males and increased estradiol and progesterone levels (peaking at copulation) in females.

High annual variability in reproductive success and survival of an Antarctic seabird, the snow petrel Pagodroma nivea : A 27-year study.

Demographic parameters were estimated for snow petrels Pagodroma nivea nesting at Pointe Géologie Archipelago, Adélie Land, Antarctica between 1963 and 1990; 21 years of data on adult survival and 27 years of data on breeding success are available. The average age of first return and first breeding were 8.1 and 9.9 years respectively and there was no signifcant difference between the sexes. The overall breeding success averaged 51.3% and was very variable between years (21-80%). Breeding failure was mostly due to incubation failure and annual breeding success was negatively correlated with average snow falls in October-November and October-March. Breeding frequency was very low, averaging 52% of seasons during a reproductive lifetime. Good quality sites, with high occupancy rate and high breeding success were few in the study plots. Poor years in 1966-1967, 1976-1977 and 1983-1984, with low breeding success, very low proportions of nets with breeding attempts and high numbers of non-breeders, occurred 1 year after large-scale El Niño Southern Oscillation (ENSO) events. Snow petrels exhibited very low philopatry. Only 45 birds have been recovered in the study plots from a total of 1115 banded fledglings giving an estimated rate of return of 12.9% between fledging and 3 years old. Annual survival between 3 and 10 years was 91.4%. Annual adult survival (93.4%), though variable, was low during poor years of 1977-1978 and 1983-1984. Adult survival of males (94.7%) was not significantly different from that of females (93.9%). Over the study period, the population of Pointe Géologie was stable. Using the estimated parameters, a Leslie model gave a growth rate of 0.948%, which was probably compensated by immigration (5.7% per year). Restricted numbers of good-quality sites at the place of birth could have led young birds to prospect other colonies and could have selected low philopatry. High adult survival, strong site tenacity and capacity to spread breeding over a long lifetime are probably part of the adaptive strategy of this small fulmarine petrel facing highly variable environmental conditions.

Over-wintering strategies of the Lesser Sheathbill Chionis minor in an impoverished and insular environment.

The Lesser Sheathbill Chionis minor is an opportunistic predator and scavenger that breeds on sub-Antarctic islands during the summer season, when many other seabirds are present, especially penguins on which it is kleptoparasitic. It is a poor flyer and over-winters on its breeding grounds, where it faces a diminished food supply and low temperatures. Its foraging behaviour was suspected to be plastic enough to respond to such a variable environment, and we show how this species enlarged its niche to survive in winter. Population movements, dietary changes and monthly weight gains were quantified during a 13-month period of observation. Mortality rates, calculated over a 19-year period of banding, appeared to be age dependent: they decrease, due to competition, from juveniles to subadults, adult non-breeders, and breeders. One of the survival strategies described here is, to our knowledge, unique among birds. After the Crested Penguin colonies have been deserted, some adult sheathbills (mainly females) remained territorial, extending their territory size and diet while their partners moved to previously undefended zones in King Penguin colonies (permanent and large in the Crozet archipelago), where they competed for a winter territory and subsequently associated with another mate. After this winter mating, mobile sheathbills returned the following summer to their breeding territory and previous mate.

Temperature regulation of young coypus (Myocastor coypus) in air and water.

Coypus (Myocastor coypus Molina) have recently colonized temperate regions of Europe in large numbers, originating from populations indigenous to subtropical regions of South America. Observations of coypu behavior in the wild and in enclosures under a temperate climate showed a preference for close proximity to water and frequent swimming bouts. Little change was observed in the semiaquatic habit in winter, despite high mortality during this period, particularly among young and adult male coypus. Evidence for constraints on thermal adaptation was sought from measurements in a thermostatic chamber of the resting metabolic rate (RMR) of young coypus caught from wild populations in France and of the body (Tb) and skin (Tsk) temperatures as a function of ambient temperature (Ta). A light heterothermic response with deep Tb falling by 0.8 degree C and stabilizing at a new level of 36.7 degrees C was observed in water and air at Ta below the lower critical temperatures. In common with other semiaquatic mammals, RMR (2.94 W/kg) was high relative to terrestrial mammals of equivalent body mass. Minimal thermal conductance was 0.10 W.kg-1.degree C-1 in air for Ta between -10 and +20 degrees C, and conductance varied in water from 0.24 to 1.18 W.kg-1.degrees C-1 for Ta between 5 and 35 degrees C. The physiological adaptations to cold and particularly the high insulative value of the fur (80-90% of the insulation) help to explain the successful extension of the coypu's range to temperate regions. Energetic costs were nevertheless high in cold water, suggesting that social factors may be implicated in severe winter mortality, particularly dominance hierarchy regulating the persistent use of water bodies even under freezing conditions.

[Electrophoretic distance between the Mandrill and the Drill (author's transl)]. / Distance électrophorétique entre le Mandrill et le Drill.

The Mandrill (Mandrillus sphinx) and the Drill (M. leucophaeus) differ by approximately 30% of their genes as shown by electrophoretic studies. The reasons of this differenciation are discussed with regard to divergence of characters, geographical isolation, and social organization of both species.