Who’s the boss? Giant petrel arrival times and interspecific interactions at a seal carcass at sub-Antarctic Marion Island

Most northern giant petrels Macronectes halli arrived before southern giant petrels Macronectes giganteus at a seal carcass at sub-Antarctic Marion Island and left earlier. In interspecific interactions, southern giant petrels initiated and dominated encounters. Level of aggression may be related to the degree of satiation. Dominance by southern giant petrels at seal carcasses may help explain why northern giant petrels commence breeding earlier than southern giant petrels in sympatric situations.     

Segregation in space and time explains the coexistence of two sympatric sub‐Antarctic petrels

Biological communities are shaped by competition between and within species. Competition is often reduced by inter‐ and intraspecific specialization on resources, such as differencet foraging areas or time, allowing similar species to coexist and potentially contributing to reproductive isolation. Here, we examine the simultaneous role of temporal and spatial foraging segregation within and between two sympatric sister species of seabirds, Northern Macronectes halli and Southern Macronectes giganteus Giant Petrels. These species show marked sexual size dimorphism and allochrony (with earlier breeding by Northern Giant Petrels) but this is the first study to test for differences in foraging behaviours and areas across the entire breeding season both between the two species and between the sexes. We tracked males and females of both species in all breeding stages at Bird Island, South Georgia, to test how foraging distribution, behaviour and habitat use vary between and within species in biological time (incubation, brood‐guard or post‐brood stages) and in absolute time (calendar date). Within each breeding stage, both species took trips of comparable duration to similar areas, but due to breeding allochrony they segregated temporally. Northern Giant Petrels had a somewhat smaller foraging range than Southern Giant Petrels, reflecting their greater exploitation of local carrion and probably contributing to their recent higher population growth. Within species, segregation was spatial, with females generally taking longer, more pelagic trips than males, although both sexes of both species showed unexpectedly plastic foraging behaviour. There was little evidence of interspecific differences in habitat use. Thus, in giant petrels, temporal segregation reduces interspecific competition and sexual segregation reduces intraspecific competition. These results demonstrate how both specialization and dynamic changes in foraging strategies at different scales underpin resource division within a community.     

Nocturnal predation of king penguins by giant petrels on the Crozet Islands

Dietary segregation of sympatric seabirds in the Southern Ocean is partly linked to differences in their foraging techniques. We have investigated the activity of giant petrels (Macronectes spp.) in a king penguin (Aptenodytes patagonicus) colony day and night during the austral winter of 2001 on the Crozet Islands. Using an automatic identification system and an infrared video camera, we followed 15 petrels tagged with micro transponders. Our data show that giant petrels predate king penguin chicks during the night. The activity of giant petrels is even slightly higher during nighttime than during the day. In addition, our data show a higher nocturnal activity by northern giant petrels (M. halli) than by southern giant petrels (M. giganteus). These unexpected results raise questions concerning visual adaptations to nocturnal foraging in giant petrels and their potential impact on the sleep, vigilance and crèching behavior of penguin chicks.     

Sexual size dimorphism in northern giant petrels: ecological correlates and scaling

Northern giant petrels ( Macronectes halli ) are among the largest and most sexually size dimorphic species of seabirds, with females being only 80% the mass of males. Both sexes scavenge on seal and penguin carrion in the sub-Antarctic ecosystem, but during the breeding season females also feed extensively on other marine food resources and show more pelagic habits than males. The outstanding sexual segregation in foraging and feeding ecology in northern giant petrels suggests that mechanisms maintaining sexual size dimorphism by ecological factors may be operating. I evaluated this possibility by examining ecological correlates with body size and by static allometry analyses. Fledging sex ratio in four consecutive years did not depart from parity. There was no assortative mating by size neither association between the male size with the breeding performance. By contrast, smaller females raised their chick in better condition. Moreover, bill size showed a size dimorphism beyond that expected by body size dimorphism, i.e. when controlling for body mass, males showed relatively longer bill than females. This trait did not deviate from isometry with respect to body size and its phenotypic variability was low, suggesting that the disproportionately large bill of males is related to their more scavenging life style compared to females. In general, the increase and maintenance of sexual size dimorphism in giant petrels is more consistent with an ecological causation rather than a result of sexual selection.     

Giant petrels as predators of albatross chicks

Giant petrels Macronectes spp. are not thought to be important predators of albatross chicks, although they are known to kill pre-fledging Thalassarche and Phoebetria albatrosses. We report the first records of predation of healthy great albatross Diomedea spp. chicks, killing wandering albatrosses D. exulans at night on sub-Antarctic Marion Island. Breeding success of this species has decreased markedly in the area where attacks occurred, suggesting that giant petrel predation events are a recent phenomenon. Mouse attacks on wandering albatross chicks may have contributed to the development of this hunting technique. We also report the first observations of giant petrel predation on pre-fledging grey-headed albatross T. chrysostoma chicks as well as additional records of sooty albatross P. fusca chicks being targeted. Only adult northern giant petrels M. halli have been confirmed to kill albatross chicks on Marion Island. Given the threatened status of wandering albatrosses, and the importance of Marion Island for this species, monitoring of their breeding success is necessary to assess whether the predation of chicks by giant petrels spreads around the island.     

Moult of the giant petrels Macronectes halli and M. giganteus at South Georgia

Moult scores were collected from colour-ringed individuals of known reproductive status of the two species of giant petrel, Macronectes halli and M. giganteus , at Bird Island, South Georgia between 1978–81.
Both species showed a substantial overlap between breeding and wing-moult, unlike most other Southern Ocean seabirds. Males started moult before females and both sexes of M. giganteus moulted at an earlier stage of the breeding cycle than M. halli , which breeds six weeks earlier than its congener.
Changes in moult rate during the breeding season are documented for both species, with Id. halli showing a rapid increase as the chick nears fledging. Male M. giganteus show a notably different pattern to the other three species-sex groups, starting moult much earlier (at egg-laying), with greater individual synchrony and usually suspending primary moult throughout the main chick growth period, whereas only two male M. halli and no females of either species suspended moult. Differences in pattern, timing and rate of moult are interpreted in terms of availability of food resources and the competing energy demands of other activities, especially chick-rearing.
Completion of primary moult could not be observed in the field but was estimated using data frcsm non-breeding birds and failed breeders; the latter started a rapid moult almost immediately they failed. In both sexes of both species moult is probably concluded at least by early winter.
The general pattern of moult in giant petrels at Bird Island is contrasted with that of other populations and species of Southern Ocean seabirds. It is suggested that the unusually extensive overlap between breeding and moult in giant petrels is a consequence of the very abundant and easily available summer food supplies (especially carrion) and the much diminished winter resources, favouring a completion of moult by the beginning of the winter.     

Origin, age, sex and breeding status of wandering albatrosses (Diomedea exulans), northern (Macronectes halli) and southern giant petrels (Macronectes giganteus) attending demersal longliners in Falkland Islands and Scotia Ridge waters, 2001–2005

A total of 547 sightings of 291 banded wandering albatrosses Diomedea exulans and 21 sightings of 14 banded giant petrels Macronectes spp. were made from toothfish longliners operating on the southern Patagonian Shelf during 2001–2005. This included 25% of the wandering albatrosses with Darvic bands that bred at Bird Island (South Georgia) during this period. Thirteen of the northern Macronectes halli and southern giant petrels Macronectes giganteus had been banded at South Georgia, and there was one sighting of a southern giant petrel from Argentina. Male and female wandering albatrosses of all age classes except young birds (<15 years old) were equally likely to attend longline vessels. Most sightings of all age classes were made during the incubation period and fewest during the brood period. Eighty-six percent of birds sighted had bred at least once before, with half currently breeding and half on sabbatical (i.e. between breeding attempts). Almost half of the wandering albatrosses were sighted on more than one occasion. The data confirms that the southern Patagonian shelf is an important foraging area for wandering albatrosses and northern and southern giant petrels, and that some individuals show consistent associations in multiple years with longline vessels fishing in the region.     

Regulation of incubation shifts near hatching by giant petrels: a timed mechanism, embryonic signalling or food availability?

In most seabird species incubation shifts shorten when hatching approaches, a behavioural response allowing the chick to be fed soon after hatching. Three mechanisms have been proposed to explain these shortened absences: endogenous timing, a response to embryo signals or a seasonal increase in food availability. I tested these hypotheses by cross-fostering eggs between two sibling species: the northern Macronectes halli and the southern M. giganteus giant petrels; the former normally breeds 5-6 weeks earlier than the latter but both have an incubation period of 60 days. The length of the last trip before hatching was unaffected by the timing of hatching, suggesting that trip length was not related to the increase in food availability with the progress of the season. Furthermore, northern giant petrels reduced their last trip lengths regardless of whether they were incubating their own developed or the southern undeveloped egg (with embryos not developed enough to signal). Parents presumably, therefore, possess endogenous control allowing them to predict hatching date and to reduce trip duration accordingly, regardless of the communication with the embryo. Nevertheless, shorter last trips before hatching in southern giant petrels incubating northern giant petrel eggs suggest that parents use embryo signals for a fine-tuning synchronization of the internal timer to the natural variability in the length of the incubation period.     

Speciation and phylogeography of giant petrels Macronectes

We examine global phylogeography of the two forms of giant petrel Macronectes spp. Although previously considered to be a single taxon, and despite debate over the status of some populations and the existence of minimal genetic data (one mitochondrial cytochrome b sequence per form), the current consensus based on morphology is that there are two species, Northern Giant Petrel M. halli and Southern Giant Petrel M. giganteus. This study examined genetic variation at cytochrome b as well as six microsatellite loci in giant petrels from 22 islands, representing most island groups at which the two species breed. Both markers support separate species status, although sequence divergence in cytochrome b was only 0.42% (corrected). Divergence was estimated to have occurred approximately 0.2 mya, but with some colonies apparently separated for longer (up to 0.5 my). Three clades were found within giant petrels, which separated approximately 0.7 mya, with the Southern Giant Petrel paraphyletic to a monophyletic Northern Giant Petrel. There was evidence of past fragmentation during the Pleistocene, with subsequent secondary contact within Southern Giant Petrels. The analysis also suggested a period of past population expansion that corresponded roughly to the timing of speciation and the separation of an ancestral giant petrel population from the fulmar Fulmarus clade.     

Hybridization and Back-Crossing in Giant Petrels (Macronectes giganteus and M. halli) at Bird Island,South Georgia,and a Summary of Hybridization in Seabirds

Hybridization in natural populations provides an opportunity to study the evolutionary processes that shape divergence and genetic isolation of species. The emergence of pre-mating barriers is often the precursor to complete reproductive isolation. However, in recently diverged species, pre-mating barriers may be incomplete, leading to hybridization between seemingly distinct taxa. Here we report results of a long-term study at Bird Island, South Georgia, of the extent of hybridization, mate fidelity, timing of breeding and breeding success in mixed and conspecific pairs of the sibling species, Macronectes halli (northern giant petrel) and M. giganteus (southern giant petrel). The proportion of mixed-species pairs varied annually from 0.4–2.4% (mean of 1.5%), and showed no linear trend with time. Mean laying date in mixed-species pairs tended to be later than in northern giant petrel, and always earlier than in southern giant petrel pairs, and their breeding success (15.6%) was lower than that of conspecific pairs. By comparison, mixed-species pairs at both Marion and Macquarie islands always failed before hatching. Histories of birds in mixed-species pairs at Bird Island were variable; some bred previously or subsequently with a conspecific partner, others subsequently with a different allospecific partner, and some mixed-species pairs remained together for multiple seasons. We also report the first verified back-crossing of a hybrid giant petrel with a female northern giant petrel. We discuss the potential causes and evolutionary consequences of hybridization and back-crossing in giant petrels and summarize the incidence of back-crossing in other seabird species.     

Sexual dimorphism,moult and body condition of seabirds killed by longline vessels around the Prince Edward Islands, 1996–97

Ryan, P.G. 1999. Sexual dimorphism, moult and body condition of seabirds killed by longline vessels around the Prince Edward Islands, 1996–97. Ostrich 70 (3&4): 187–192.

A total of 393 seabirds from nine species killed by longline fishing in South African waters around the Prince Edward Islands during summer 1996–97 were sexed, measured, and their body fat levels and primary moults scored. Males of Whitechinned Petrels Procellaria aequinoctialis and Greyheaded and (Indian Ocean) Yellownosed Mollymawks Thalassarche chrysostoma and T. [chlororhynchosj bassi averaged 1–9% larger than females. Measures of bill depth were the most dimorphic characters in all three species. I recommend that if only a single measure is taken to estimate sex of adults, bill depth at the base be the standard measure for petrels, and bill depth at the nail be the standard measure for mollymawks. Discriminant functions are described for predicting the sex of unknown individuals; the proportion of misclassified birds ranged from 2–10%. With the exception of giant petrels Macronectes spp., only two young birds were actively moulting their primary flight feathers. Analysis of wear patterns among adult Greyheaded Mollymawk primaries conforms with studies of moult in this species at South Georgia. Fat scores did not differ between sexes. They tended to decrease during the breeding season, but trends were masked by great inter-individual variation. There was no significant relationship between fat scores and the presence of ingested plastics. Plastic debris was found in the stomachs of 90 birds from five species. Incidence in Whitechinned Petrel stomachs (37%) was less than that reported from birds collected off South Africa in the early 1980s (57%), but this is probably due to the predominance of breeding adults. The frequency and size of plastic loads decreased through the breeding season, which is consistent with the hypothesis that inter-generational transfer is important in the dynamics of ingested debris.     

Food, feeding ecology and ecological segregation of seabirds at South Georgia

At the sub-Antarctic island of South Georgia 25 of the 29 breeding species are seabirds. Fifteen of these have recently been studied in some detail. By examining the timing of their breeding seasons and their diet and feeding ecology (especially feeding techniques and potential foraging ranges), the nature of their ecological isolating mechanisms, and in particular the way in which they partition the resources of the marine environment, are reviewed.
Although breeding season adaptations occur (winter breeding in Wandering Albatross and King Penguin; out of phase breeding in two species-pairs of small petrels) these are less important than dillerences in food and feeding ecology. There is a fundamental distinction between the niche of pursuit-diving species (mainly penguins) and the remainder which are basically surface-feeders. The two abundant krill-eating penguins show clear differences in feeding zones. Three albatrosses and a petrel feed mainly on squid and there are differences in both the species and size of the prey of each. The remaining seabirds chiefly take krill (although the giant petrels are extensive scavengers and some smaller petrels specialize on copepods) and utilize different feeding methods and areas to do so.
Various adaptations related to inshore and offshore feeding zones are discussed. Although most species possess a combination of ecological isolating mechanisms additional evidence for the particular importance of dietary differences is presented.     

Sexual dimorphism in weight among the primates: The relative impact of allometry and sexual selection

In this paper, we examine allometric and sexual-selection explanations for interspecific differences in the amount of sexual dimorphism among 60 primate species. Based on evidence provided by statistical analyses, we reject Leutenegger and Cheverud’s [(1982). Int. J. Primatol.3:387-402] claim that body size alone is the major factor in the evolution of sexual dimorphism. The alternative proposed here is that sexual selection due to differences in the reproductive potential of males and females is the primary cause of sexual dimorphism. In addition, we propose that the overall size of a species determines whether the dimorphism will be expressed as size dimorphism,rather than in some other form.     

Sex‐specific patterns of fuelling and pre‐breeding body moult of Little Stints Calidris minuta in South Africa

The timing and duration of each stage of the life of a long‐distance migrant bird are constrained by time and resources. If the parental roles of males and females differ, the timing of other life stages, such as moult or pre‐migratory fuelling, may also differ between the sexes. Little is known about sexual differences for species with weak sexual dimorphism, but DNA‐sexing enables fresh insights. The Little Stint Calidris minuta is a monomorphic long‐distance migrant wader breeding in the Arctic tundra. Males compete for territories and perform elaborate aerial displays. Females produce two clutches a season. Each sex may be a bigamist and incubate one nest a season, each with a different partner. We expect that these differences in breeding behaviour entail different preparations for breeding by males and females, so we aimed to determine whether Little Stints showed any sex differences in their strategies for pre‐breeding moult and pre‐migratory fuelling at their non‐breeding grounds in South Africa. We used body moult records, wing length and body mass of 241 DNA‐sexed Little Stints that we caught and ringed between 27 January and 29 April in 2008–2018 at two neighbouring wetlands in North West Province, South Africa. For each individual we assessed the percentage of breeding plumage on its upperparts and took blood samples for DNA‐sexing. We calculated an adjusted Body Moult Index and an adjusted Wing Coverts Moult Index, then used the Underhill–Zucchini moult model to estimate the start dates and the rate of body moult in males and females. We estimated the changes in the sex ratio of the local population during their stay in South Africa, and also estimated the timing and rate of pre‐migratory fuelling and the potential flight ranges for males and females. The males started body moult on average on 7 February and the females on 12 February, but the sexes did not differ in their timing of wing covert moult, which started on average on 10 February. In January to mid‐February, males constituted c. 57% of the population, but their proportion declined afterwards, indicating an earlier departure than females. We estimated that both sexes began pre‐migratory fuelling on average on 15 March. The sexes did not differ in fuelling rate, but most females stayed at the non‐breeding site longer than the males, and thus accumulated more fuel and had longer potential flight ranges. These patterns of moult and fuelling suggest sex differences in preparations for breeding. We suggest that the males depart from South Africa earlier but with smaller fuel loads than the females to establish breeding territories before the females arrive. We conclude that for each sex the observed trade‐offs between fuelling and moult at the non‐breeding grounds are precursors to different migration strategies, which in turn are adaptations for their different roles in reproductive behaviour.     

Insights into the genetic architecture of sexual dimorphism from an interspecific cross between two diverging Silene (Caryophyllaceae) species

The evolution of sexual dimorphism in species with separate sexes is influenced by the resolution of sexual conflicts creating sex differences through genetic linkage or sex‐biased expression. Plants with different degrees of sexual dimorphism are thus ideal to study the genetic basis of sexual dimorphism. In this study we explore the genetic architecture of sexual dimorphism between Silene latifolia and Silene dioica. These species have chromosomal sex determination and differ in the extent of sexual dimorphism. To test whether QTL for sexually dimorphic traits have accumulated on the sex chromosomes and to quantify their contribution to species differences, we create a linkage map and performed QTL analysis of life history, flower and vegetative traits using an unidirectional interspecific F2 hybrid cross. We found support for an accumulation of QTL on the sex chromosomes and that sex differences explained a large proportion of the variance between species, suggesting that both natural and sexual selection contributed to species divergence. Sexually dimorphic traits that also differed between species displayed transgressive segregation. We observed a reversal in sexual dimorphism in the F2 population, where males tended to be larger than females, indicating that sexual dimorphism is constrained within populations but not in recombinant hybrids. This study contributes to the understanding of the genetic basis of sexual dimorphism and its evolution in Silene.     

Factors influencing the degree of sexual size dimorphism within and among calanoid copepod species

Populations of Diaptomus leptopus (Copepoda: Calanoida) and other calanoid copepods exhibit varying degrees of sexual size dimorphism. We examined whether intraspecific or interspecific variation in dimorphism could be explained by allometry, and we examined the relationship between adult size attained and development rate to determine any relationship between the two. We compared the degree of sexual size dimorphism in D. leptopus and in other calanoid copepods inhabiting temporary and permanent habitats. Allometry did not explain variation in sexual size dimorphism within or among populations or among species. Permanence of habitat affected the degree of dimorphism: dimorphism was greater within and among species inhabiting temporary environments. Non-significant differences in development rate were found among populations and significant differences were found between sexes of D. leptopus when reared under identical laboratory conditions: males developed more rapidly than females but there was no general relationship between development rate and adult size. Potential adaptive hypotheses to explain the differences between populations inhabiting temporary and permanent habitats are discussed.     

Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae,Caimaninae)

Melanosuchus niger is a caimanine alligatorid widely distributed in the northern region of South America. This species has been the focus of several ecological, genetic and morphological studies. However, morphological studies have generally been limited to examination of interspecific variation among extant species of South American crocodylians. Here, we present the first study of intraspecific variation in the skull of M. niger using a two‐dimensional geometric morphometric approach. The crania of 52 sexed individuals varying in size were analysed to quantify shape variation and to assign observed shape changes to different types of intraspecific variation, that is, ontogenetic variation and sexual dimorphism. Most of the variation in this species is ontogenetic variation in snout length, skull depth, orbit size and the width of the postorbital region. These changes are correlated with bite force performance and probably dietary changes. However, a comparison with previous functional studies reveals that functional adaptations during ontogeny seem to be primarily restricted to the postrostral region, whereas rostral shape changes are more related to dietary shifts. Furthermore, the skulls of M. niger exhibit a sexual dimorphism, which is primarily size‐related. The presence of non‐size‐related sexual dimorphism has to be tested in future examinations.     

Population aspects of sexual dimorphism in Mustelidae from the example of four species (Mustela lutreola, Neovison vison, Mustela putorius, and Martes martes)

Size sexual dimorphism was studied on 695 crania of four species of the Mustelidae. According to the degree of increasing differences between sexes, the species were arranged in the following order: common marten (Martes martes), European mink (Mustela lutreola), American mink (Neovison vison), and black polecat (Mustela putorius). The value of dimorphism characterizes the ecological flexibility of species and is an integrated population characteristic. It was demonstrated that M. martes occupies a specific and relatively narrow ecological niche of the forest ecosystem and joins weak competitive relations with smaller species of Mustelidae. The level of sexual dimorphism in M. lutreola and N. vison, as well as M. putorius, reflects the tension of their interspecific relationships in a given territory. A high level of sexual dimorphism in M. putorius is caused by further divergence of the ecological niches of males and females, being a compensatory mechanism, which mitigates the consequences of toughened environmental requirements.     

Metric variability in the anterior dentition of African colobines

The anterior dentition of three species of African colobines (Colobus polykomos, C. badius, and C. verus) was investigated metrically and the results analyzed for three characters: (1) intraspecific tooth size relations, (2) sexual dimorphism, and (3) interspecific relations. Based on incisor size sequences C. polykomos and C. badius appear to be more closely related to each other than either is to C. verus. However, incorporating the results of a previous study on postcanine dentition the three species appear to be equally closely related. The magnitude of sexual dimorphism in canine size decreases from C. badius to C. verus to C. polykomos. Interspecific differences in the degree of canine size dimorphism may be attributed to differential intensities of male intrasexual selection; however, the interspecific differences in canine size dimorphism do not correspond to the interspecific differences in body size dimorphism.     

The timing of gonadal development and moult in three raptors with different breeding seasons: effects of gender, age and body condition

Differences between species in breeding seasons are thought to be mediated through differences in their reproductive physiology. Little is known about how the timing and duration of gonadal maturation varies between raptor species, how the timing of moult relates to the gonadal cycle, whether the timing and degree of sexual maturation varies between juveniles and adults or whether body condition has a significant effect. To address these questions, data on gonadal size and moult for adults and juveniles of both sexes of three raptor species were extracted from the Predatory Bird Monitoring Scheme (based on birds found dead by members of the public). The three species, Sparrowhawk Accipiter nisus, Kestrel Falco tinnunculus and Barn Owl Tyto alba, have different ecologies – diurnal bird predator, diurnal mammal predator and nocturnal mammal predator, respectively. All are single‐brooded but have different breeding seasons. The duration of gonadal maturation was markedly different between the species. Barn Owls showed the earliest maturation and the latest gonad regression, and Sparrowhawks the latest maturation and earliest gonad regression. Kestrels were intermediate. In males of all species, the testes remained fully mature throughout their respective breeding seasons. In females, the ovaries remained partially mature throughout the breeding season. Moult started slightly earlier in Sparrowhawks than in Kestrels and coincided with gonadal regression in the two species. Although females of the two species started to moult earlier than males, moult duration was similar between the sexes. Barn Owls showed no distinct annual pattern of moult. In juveniles of all three species, the gonads were smaller than in adults throughout spring and started to mature later. Gonad size in birds that had starved tended to be smaller than in birds dying from other causes, but did not influence the difference in gonad mass between adults and juveniles and between seasons. Body condition had no effect on moult. Whilst ecology has led to the evolution of different breeding seasons, differences between species, and between adults and juveniles, are mediated through adaptive differences in their reproductive physiology.