Behavioral and ecological interactions of foraging mice (Peromyscus melanotis) with overwintering monarch butterflies (Danaus plexippus) in México

Summary Mice (Peromyscus melanotis) immigrate extensively to overwintering colonies of monarch butterflies (Danaus plexippus) in México. There they feed on both live and dead butterflies that accumulate on the ground and in low vegetation. Through a series of feeding experiments, we examined the potential impact of mouse predation on these colonies, as well as how this predation was influenced by the accessibility and the degree of desiccation of the monarchs. Mice attacked on average 39.9 wet (freshlykilled) butterflies per night. We estimated that a population of mice (75–105 individuals) could kill approximately 0.40–0.57 million butterflies in a 1 ha colony (4–5.7% of the colony) over the 135-day overwintering season. In feeding experiments, mice fed disproportionately on: 1) wet (hydrated) monarchs close to the ground versus those perched higher; 2) wet monarchs, when both wet and dry (desiccated) monarchs were on the ground; and 3) wet monarchs on stakes versus dry monarchs on the ground. Mice commonly ate the entire abdomen of dry monarchs, whereas they fed selectively on the abdomen of wet monarchs by discarding the bitter, cardenolide-laden cuticle and eating the internal tissues. These results suggest that the monarchs' state of desiccation is more important than their accessibility in determining the feeding preferences of these mice. However, the monarchs' strong tendency to crawl up vegetation does appear to reduce their risk to mouse predation.     

Overwintering ecology of the monarch butterfly, Danaus plexippus L., in California

Abstract. 1. Population and physiological measurements were made on monarch butterflies in three overwintering colonies along the California coast during the 1975–76 season. 2. Mark, release and recapture studies indicated that two northern colonies (Muir Beach and Santa Cruz) stabilized at maxima of about 40000 and 95 000 individuals with little movement in or out of colonies, whereas the southern one at Santa Barbara appeared dynamic with a maximum of about 45 000. 3. Initially low, sexual activity builds to a massive mating ceremony in February, followed by rapid colony dispersal. The butterflies deteriorate gradually during the overwintering period, but males do so to a greater extent because of their involvement in frenzied courting flights. 4. Changes in several parameters at the Muir Beach colony were measured between 1 November and 2 February. Right forewing length did not change with time, indicating no differential mortality with respect to size. However, both sexes lost about 25% of their total dry weight, and 49–60% of their lipid contents. Minimum fat levels were about 20 mg and some butterflies were as lean on arrival as were the most depleted overwintering survivors. 5. The importance of nectaring in restoring lipid reserves is potentially great but remains uninvestigated. The hypothesis is put forward that the distance spring migrants are capable of flying is dependent largely upon their lipid reserves following the overwintering period. This and the intracolony mass-mating ceremony are seen as randomizing factors which prevent geographic differentiation and allow the maintenance of the generalized, nature of the migration, which in turn enables the monarch to exploit the vast milkweed flora across the North American continent. 6. Cardenolides also diminished during the wintering, but to a substantially lesser extent than the lipids. There was no evidence for a change in the proportion of cardenolide-positive butterflies, suggesting no differential elimination by predation of the poison-free palatable individuals. This is the first field evidence in support of the theory of automimicry. 7. The overwintering behaviour of the monarch butterfly is seen as an integrative force in optimizing foodplant exploitation, reproduction, anti-predator, and migration strategies.     

Energy reserves and metabolic expenditures of monarch butterflies overwintering in southern California

Abstract. 1. Energetic expenditure and predicted requirements for overwintering metabolism were determined for monarch butterflies ( Danaus plexippus L.) in southern California.
2. Fat content of butterflies declined steadily from a maximum of 71% lean dry weight in late November to a minimum of 36% lean dry weight in late January. The energy expenditure measured by fat depletion over a 61 day period from 24 November to 25 January was 26.05 joules per day.
3. Butterflies were generally the same temperature as the air at any time they were inactive, whether they were part of a large cluster or roosting solitarily.
4. Oxygen consumption of butterflies increased in a curvilinear fashion with increasing air temperature. Thus, the predicted metabolic requirements for an inactive butterfly during their quiescent period from late November to late January was 30.32 joules per day.
5. In contrast to the steady depletion of fat reserves during their quiescent period in December and January, low and stable fat levels of butterflies in late February coincide with high levels of flight activity, mating and emigration of females from the colony.
6. Progressive tightening of the energy balance due to increasing nocturnal temperatures from northern to southern California coastal areas may limit the southern distribution and duration of overwintering aggregations.     

Sexual and seasonal differences in the frequency of beak marks on the wings of two Lethe butterflies

To assess bird predation pressure on butterflies, I investigated beak marks on the wings of two Lethe butterflies for 3 years in secondary temperate forests. If bird predation had significant effects on average longevity of butterflies, and if the number of specimens preyed upon was proportionate to the number of beak-marked specimens, the beak mark frequency would be negatively correlated with average longevity of a butterfly. Bird predation pressure is generally thought to influence average longevity of butterflies. Therefore, if there is a negative correlation between beak mark frequency and average longevity, bird predation pressure would be reflected in beak mark frequency. Beak mark frequency was negatively correlated with longevity in Lethe diana (Butler), the more abundant of the two species; thus, the beak mark frequency was considered to be a suitable index of bird predation pressure on the butterflies investigated in this study. In both Lethe species, beak mark frequency was higher in females than in males. Because female butterflies have a relatively smaller thorax and flight muscles and a larger abdomen that contains eggs, they are presumably weaker or less agile fliers than males, and are probably attacked more easily by birds. In autumn, butterflies were heavily attacked by birds irrespective of sex and species. Because the numbers of lepidopteran larvae, which are the preferred prey of many birds, decreased in autumn, birds were thought to shift their diets to alternative prey such as adult butterflies.     

New defensive chemical data for ground beetles (Goleoptera: Carabidae): interpretations in a phylogenetic framework

First reports of major defensive chemicals for ground beetles representing four tribes include: Morionini (formic acid), Dercylini (methacrylic and tiglic acids), Catapieseini (formic acid and decyl acetate) and Perigonini (formic acid and dccyl acetate). Multiple species from Loxandrini were sampled and, shown to contain formic acid, not salicylaldehyde as previously reported. Several hexenoic acid compounds were found in the clivinine genus Schizogenius representing a third class of chemicals for that tribe. Salicylaldehyde was found for the first time in a species of Oodini. Additional species from Pterostichini, Patrobini and Odacanthini were sampled and the results were found to be consistent with previously published reports. The taxonomic distribution of defensive secretions is reviewed for tribes across the family Carabidae. The simultaneous occurrence of hydrocarbons and formic acid is noted in phylogenetically more derived carabids. By mapping chemical classes onto a phylogenetic hypothesis, it is shown that formic acid or other relatively strong irritants are correlated with tribes having a high species diversity in tropical regions, whereas tribes exhibiting higher diversity in temperate regions use milder saturated/unsaturatcd earboxylic acids. Based on this phylogenetic interpretation, the evolution and maintenance of formic acid is interpreted as the result of predation pressures and possibly the evolution of chemical mimicry.     

Sign left by introduced and native predators feeding on Hutton's shearwaters Puffinus huttoni

Abstract

The identification of introduced and native predators is important for many conservation studies within New Zealand. Carcasses of Hutton's shearwaters were collected over three field seasons, and where predation was probable, the bodies were autopsied. Paired bites identified stoats as the principal predator of Hutton's shearwater, but also revealed that a feral cat was present within the colony. Stoats killed their prey with a bite to the back of the neck or head, and commenced feeding on the neck or head. Despite the limited number of cat‐killed birds, cats appeared to feed on Hutton's shearwaters differently from stoats, starting on the breast muscles. Harriers and kea left sign that allowed birds killed or scavenged by these native birds to be distinguished from those killed by stoats or cats.     

Ringing or colour-banding does not increase predation mortality in redshanksTringa totanus

The use of metal and colour‐rings or bands as a means of measuring survival, movements and behaviour in birds is universal and fundamental to testing ecological and evolutionary theories. The practice rests on the largely untested assumption that the rings do not affect survival. However this assumption may not hold for several reasons, for example because the ‘oddity effect’ predicts predators select prey that appear different to their neighbours in order to avoid the ‘confusion effect’. We compared the foraging behaviour and the death rates of redshanks Tringa totanus conspicuously marked with six colour rings and one metal ring each to unmarked birds in a study system, where routinely up to 50% of the total population are killed by avian predators during a winter. If avian predators selectively target and/or have a higher capture success of ringed birds then we would predict the proportion of colour‐ringed birds in the population to decline through a winter. The proportion of colour‐ringed birds in the population did not change over the course of three separate winters, and in one winter the ratio of marked:unmarked birds found killed by sparrowhawks Accipiter nisus was the same as the ratio of marked birds alive in the population. In the year with largest sample size, power was sufficient to detect a greater than 2.2% difference in predation rate between ringed and unringed groups. The average kill rate difference between ringed and unringed birds across the three winters was less than 1% (0.73±2.2%) suggesting that even if there were differences in predation rate that were not detected because of low statistical power they were extremely small. There were no differences in any foraging measures comparing ringed and unringed birds, suggesting that the rings did not affect the ability of birds to meet their daily energy budgets. The results showed that colour‐ringed birds were not preferentially targeted or killed by avian predators, and suggest that the presence of a metal and even several large colour‐rings is unlikely to affect behaviour and predation mortality even under extreme selection.     

Long‐term and large‐scale analyses of nest predation patterns in Australian songbirds and a global comparison of nest predation rates

Juvenile mortality is one of crucial drivers of life‐history evolution, and predation is the main cause of nest loss in birds. Thus, understanding how nest predation and failure vary in nature is important for understanding life history evolution and, moreover, for effective conservation. We used published data and unpublished records to study factors influencing nest predation and total failure in 138 populations of 90 species of Australian songbirds. Daily predation (average 2.0% d^?1) and failure rates (2.9%) increased from temperate regions to the tropics, over the last four decades, and were lowest in temperate south‐western Australia. Predation and failure were higher in smaller species, and failure rates were lower in species with closed nests than in species with open nests. There was no effect of nest height or nest site (ground, shrub, canopy) or social organization on nest predation or failure rates. Nest predation caused on average 72% of total nest failure, similar to other tropical, subtropical, and temperate areas. Our study spanning from the tropics to temperate regions and using > 10 000 nests confirmed that tropical birds faced higher nest failure rates. We identified an increase in nest depredation rates in the last four decades in Australia, suggesting that a large‐scale ecological phenomenon must be responsible. It may include increases in predator abundances and/or ranges, possibly connected with human‐caused habitat change. A global comparison of nest failure rates confirmed that predation is the main source of nest mortality in songbirds worldwide. We discuss implications of our results for the evolution of reproductive strategies and for the conservation of Australian birds.     

Palatablility and escaping ability in Neotropical butterflies: tests with wild kingbirds (Tyrannus melancholicus, Tyrannidae)

The palatability and the ability of neotropical butterflies to escape after being detected, attacked and captured by wild kingbirds ( Tyrannus melancholicus Vieillot), was investigated by the release of 668 individuals of 98 butterfly species close to the birds, during their usual feeding activities. Most of the butterflies were attacked and eaten. Only the troidine swallowtails ( Parities and Battus ; Papilionidae) were consistently rejected on taste and elicited aversive behaviours in birds. Most other aposematic and/or mimetic species in the gehera Danaus and Lycorea (Danainae), Dione, Eueides and Heliconius (Heliconiinae), Hypothyris, Mechanitis and Melinaea (Ithomfinae), Biblis, Callicore and Diaethria (Limenitidinae) were generally eaten. Cryptic and non-mimetic species were always attacked and, if captured, they were also eaten. All Apaturinae, Charaxinae, Nymphalinae, Hesperidae, most Limenitidinae, Heliconiinag ( Agraulis, Dryas, Dryadula and Philaethria ) and Papilionidae ( Eurytides, Heraclides and Protesilaus ) were in this group. Results indicate that the learning process in kingbirds may demand a large mortality in prey populations, even among species generally accepted as unpalatable and aposematic. They also support the assertion that escaping ability and unpalatability evolved in butterflies as alternative strategies to avoid predation by birds. Mimetic relationships among several species are discussed. Evidence for the evolution of aposematism not related to unpalatability, but to escaping ability, was found for two hard-to-catch Morpho species.     

Safety first: terrestrial predators drive selection of highly specific nesting sites in colonial‐breeding birds

Nesting is a critical yet hazardous life stage for many birds. For colonial‐breeding birds, the conspicuousness of the colony to predators suggests immense pressure to select optimal colonial nesting sites. But what drives selection of those sites? As with solitary nesting birds, reducing access by predators may be the single most important factor. If so, knowledge of the predators involved and the attributes of different potential colony sites can allow us to predict the features that make a site especially safe. We examined the attributes of trees used by breeding colonies of metallic starlings Aplonis metallica in tropical Australia, and experimentally tested if those attributes prevented nest access by predatory snakes. Our surveys confirmed that tree choice by starling nesting colonies is highly non‐random, with all colonies located in tall trees in rainforest clearings, with no low branches and smooth bark. Experimental tests demonstrated that the climbing ability of predatory snakes depends upon bark rugosity, and that colony access by snakes depends on tree attributes such as bark rugosity and canopy connectivity. Our study confirms that colonial‐nesting starlings select colony sites that provide a safe refuge from predation. Intense predation pressure may have driven the evolution of stringent breeding habitat criteria in many other species of colonial‐breeding birds.     

The starvation–predation risk trade-off, body mass and population status in the Common Starling Sturnus vulgaris

It is theoretically and empirically well established that body mass variation in small birds reflects a trade-off between starvation risk and predation risk. This occurs because carrying increased fat reserves reduces starvation risk but also results in a higher predation risk due to reduced escape flight performance and/or the increased foraging exposure needed to maintain a higher body mass. In principle, therefore, the theory of mass-dependent predation risk could be used to understand how a bird perceives and responds to the risks in its environment, because its mass will reflect the predictability of foraging opportunities and predation risk. Mass in birds may then provide a relatively straightforward way of assessing the foraging environment of birds and so the potential conservation problems a species faces. This study tests, for the first time for any species, how body mass changes in response to changing starvation risk, changing predation risk and changing population status. Common Starling Sturnus vulgaris mass varies as predicted by starvation–predation risk trade-off theory: mass is lower when foraging conditions are more favourable and when predation risk is increased. The populations that are declining the most strongly have higher mass, which is most likely indicative of a poor foraging environment, leading to lower relative survival. The results suggest that increased mass in Starlings, and possibly in other species, may provide an indication of the poor quality of the foraging environment and/or rapidly declining populations.     

BREEDING PERFORMANCE OF PUFFINS FRATERCULA ARCTICA IN RELATION TO NEST DENSITY, LAYING DATE AND YEAR

The paper presents data on the breeding and predation of Puffins in two areas of different nest density within a single colony on Dun, St Kilda group, Outer Hebrides in 1973-78.
Within a season birds laying early had a slightly higher nesting success than birds laying late, but laying date had little influence on the peak and fledging weights of young. The main disadvantage in late laying was a reduced chance of relaying if the first egg was lost.
Breeding success and chick weights varied from year to year. The 1974 season was the least successful with the lowest nesting success, lowest frequency of feeds, lowest calorific value of feeds, lightest chicks and slowest growth. Overall breeding performance was not related to the annual mean laying dates.
In all years pairs nesting in the area of high nest density did better than pairs nesting at low density. The effect is attributed to differential predation and disturbance by predatory gulls. At least 4.2% of adult Puffins breeding in the area of low burrow density were killed by gulls each breeding season; this is higher than the total annual mortality found in three other studies. Only 0.9% of adults from the high density area were found killed. The subpopulation in the low density area cannot survive without much immigration, yet there is no evidence that this happens.     

Colony size selection determines adult survival and dispersal preferences: allee effects in a colonial bird

Avian coloniality traditionally has been investigated by examining how breeding success varies with colony size, but other crucial fitness components rarely have been examined. This may lead to wrong conclusions because unmeasured parameters may change the final fitness balance. We used multistate capture-recapture models to investigate adult survival and dispersal in relation to colony size within a long-term monitored population of lesser kestrels (Falco naumanni). Nest predation probability decreases with colony size, and adult survival is predicted to show the same trend because adults are exposed to the same suite of predators. As expected, survival probability was higher in large colonies (0.72+/-0.015; mean+/-SE) than in medium or small colonies (0.65+/-0.02). Additionally, dispersal probabilities were higher going from small to large colonies (0.20+/-0.01) than from large to small (0.08+/-0.01), as predicted by theory of habitat selection shaped by fitness maximization. These asymmetries are likely to generate size-specific colony population dynamics, so they should be taken into account in studies of colonial birds and other metapopulation-like systems. Allee effects, that is, positive density dependence, appear to be the cause of the evolution of dispersal behavior and may explain the maintenance of coloniality in this species.     

Causes of morbidity and mortality and their effect on reproductive success in double-crested cormorants from Saskatchewan

The objectives of this study were to describe causes of morbidity and mortality in a breeding colony of double-crested cormorants (Phalacrocorax auritus) on Doré Lake (Saskatchewan, Canada), and to determine cause-specific mortality rates of juvenile birds. Morbidity and mortality were monitored every third day during the breeding season from 1994 to 1996 from inside a tunnel-and-blind system. Affected eggs and birds were collected for examination and diagnosis. The cause of mortality was determined for 105 eggs, 178 nestlings (< or = 4-wk-old), 1393 post-nestling chicks (> 4-wk-old), and 10 adults. The main causes of mortality were infertility or embryonal death, avian predation, displacement of eggs and chicks from the nest, starvation from sibling competition, Newcastle disease, coyote predation, human-induced suffocation, and entrapment. In 49% of the cases, avian predation and displacement from the nest of eggs or nestlings was associated with human disturbance. Thirty-six nestlings, 40 post-nestling chicks, and three adults were examined for the presence of parasites. Contracaecum spiculigerum was found in the proventriculus; Amphimerus elongatus in the liver. Piagetiella incomposita in the gular pouch; Eidmanniella pellucida, Pectinopygus farallonii, and Ceratophyllus lari in the plumage; and Theromyzon sp. in the nasal and oral cavity. Contracaecum spiculigerum was associated with ulcerative gastritis, A. elongatus with multifocal hepatitis and bile duct hyperplasia, and P. incomposita with ulcerative stomatitis, but these lesions were not considered fatal. Other diseases included beak deformity, abnormal rotation of the carpal joint, hypopigmentation, and eye loss. Overall mortality of cormorant chicks between hatching and the end of the breeding season varied from 25 to 48%. The most important causes of mortality were Newcastle disease, which killed 21% of hatched chicks in 1995, sibling competition (maximum 12% in 1994), and coyote predation (2% in 1994).     

Notes on New Zealand mammals 8. Predation on nesting southern royal albatrosses Diomedea epomophora by a New Zealand sea lion Phocarctos hookeri

Abstract

This note documents unusual predation of southern royal albatrosses Diomedea epomophora on land by an individual New Zealand sea lion Phocarctos hookeri. During a survey of albatross nests on Campbell Island in the New Zealand sub‐antarctic in January 2005, 128 corpses of adult albatrosses were found at or near nests within a 230 ha area. Flattened vegetation and flipper prints near the nests suggested sea lion predation, and subsequently a male sea lion was found near a line of freshly killed birds. The predation rate was estimated to be at least seven birds per day over a number of weeks; similar skeletal remains from the previous year were observed in a neighbouring area. The impact of one threatened species on another can be a dilemma for conservation managers. In this case, apparently only one male sea lion was involved and, because of the ongoing impact to the albatross population, permission was obtained to cull that individual on 17 January 2005. No further incidents were observed over the following three seasons (2006–08).     

Freeze-protection of overwintering monarch butterflies in Mexico: critical role of the forest as a blanket and an umbrella

Abstract. 1. At their high-altitude overwintering sites in Mexico, monarch butterflies frequently are subjected to sub-zero°C temperatures during December-March. Although monarchs have moderate supercooling ability, two ecological factors strongly influence their capacity to resist freezing: wetting and exposure to the clear night sky. 2. As shown in Fig. 2, 50% of a population of butterflies with water on their body surfaces freeze at warmer sub-zero temperatures (-4.2°C) compared to butterflies with no water on their bodies (-7.7°C). 100% mortality occurs, respectively, at ?7.7°C and ?15°C. 3. Comparative measurements of rainfall within a large overwintering colony in Mexico indicated that the intact canopy acts as an umbrella that reduces butterfly wetting during winter storms. 4. Variable experimental exposure of butterflies to the clear night sky indicated that openings in the forest canopy increases radiational cooling and causes monarch body temperatures to drop as much as 4°C below ambient air temperature. Monarchs under dense cover had body temperatures approximately the same as the ambient air temperature, but more exposed individuals had body temperatures below ambient in direct proportion to the degree of exposure. Consequently, forest thinning increases the probability that the butterflies will freeze to death. 5. Whereas both wetting and exposure are increased by disturbance of the forest canopy, the interaction of these two factors exacerbates freezing mortality during winter storms: 50% of dry and unexposed butterflies froze at ?8°C, whereas wetted and fully exposed butterflies froze at only ?0.5°C. 6. Butterflies inside and on the bottom of the fir bough clusters are better protected from wetting than those on the outside. This supports the hypothesis that the structure of the butterfly clusters has evolved through individual selection to avoid wetting. 7. The data strongly reinforce previous evidence that forest thinning should be totally prevented within and adjacent to the overwintering sites in order to minimize both wetting and exposure of the butterflies that synergistically increase winter mortality at the overwintering sites in Mexico.     

Factors favouring the development and maintenance of outbreaks in an invasive leaf miner Cameraria ohridella (Lepidoptera: Gracillariidae): a life table study

1 Life tables were constructed for seven generations of the invasive horse‐chestnut leaf miner Cameraria ohridella in Switzerland, to assess the factors allowing the moth to build and maintain outbreak densities and to identify ‘gaps’ among their mortality factors that could be targeted by new control methods. The fecundity of females was calculated and the mortality factors affecting all developmental stages determined. 2 Females contained 106.6 eggs. In captivity, the mean number of eggs laid per female varied between 34 and 82 eggs, with a maximum of 184 eggs. Egg mortality was 18–75% and was mostly due to unknown factors. 3 The main mortality factors affecting larvae and pupae in the leaves were predation by birds and arthropods in all generation and leaf senescence in the autumn generation. Parasitoids killed less than 5% of the population. 4 Nearly 90% of overwintering pupae died in dead leaves, the majority of them being killed by earthworms and other leaf decomposers. As a result, the overwintering generations were the only ones showing a net rate of increase less than one. Non‐overwintering generations had net rates of increase between four and 11, allowing populations to build up from spring to autumn. 5 The persistence of high outbreak densities in Europe is explained by a combination of at least three factors: (i) high fecundity; (ii) multivoltinism; (iii) the unusually low impact of traditional natural enemies of leaf miners and particularly parasitoids. 6 Yearly variations in population densities could be at least partly explained by differences in larval mortality due to leaf senescence and intraspecific competition in the last generation in autumn.     

Cryptic differences in colour among Müllerian mimics: how can the visual capacities of predators and prey shape the evolution of wing colours?

Antagonistic interactions between predators and prey often lead to co‐evolution. In the case of toxic prey, aposematic colours act as warning signals for predators and play a protective role. Evolutionary convergence in colour patterns among toxic prey evolves due to positive density‐dependent selection and the benefits of mutual resemblance in spreading the mortality cost of educating predators over a larger prey assemblage. Comimetic species evolve highly similar colour patterns, but such convergence may interfere with intraspecific signalling and recognition in the prey community, especially for species involved in polymorphic mimicry. Using spectrophotometry measures, we investigated the variation in wing coloration among comimetic butterflies from distantly related lineages. We focused on seven morphs of the polymorphic species Heliconius numata and the seven corresponding comimetic species from the genus Melinaea. Significant differences in the yellow, orange and black patches of the wing were detected between genera. Perceptions of these cryptic differences by bird and butterfly observers were then estimated using models of animal vision based on physiological data. Our results showed that the most strikingly perceived differences were obtained for the contrast of yellow against a black background. The capacity to discriminate between comimetic genera based on this colour contrast was also evaluated to be higher for butterflies than for birds, suggesting that this variation in colour, likely undetectable to birds, might be used by butterflies for distinguishing mating partners without losing the benefits of mimicry. The evolution of wing colour in mimetic butterflies might thus be shaped by the opposite selective pressures exerted by predation and species recognition.     

Flight morphology of Neotropical butterflies: palatability and distribution of mass to the thorax and abdomen

Summary We test whether palatability of Neotropical butterflies is associated with distribution of mass to the thorax and abdomen. Thoracic mass is predominantly muscle mass, whereas abdominal mass includes organs of digestion, food storage, and reproduction. To escape from predation, butterflies palatable to the rufous-tailed jacamar (Galbula ruficauda) use fast, erratic flight, whereas unpalatable butterflies have defensive chemicals and slow, regular flight patterns. We adjusted for effects of phylogeny and report partial correlations for two levels of analysis: 1) comparisons among-lineage means, which test for correlations between traits of distantly related lineages, and 2) comparisons among deviations from lineage means (or within lineages), which test for correlations between traits of more closely related species.Among lineages for both males (n=10 lineages) and females (n=9), palatability and thoracic mass were positively correlated, whereas palatability and abdominal mass were negatively correlated. An inverse correlation between thoracic and abdominal mass is a consequence of the two segments composing 75% of the total body mass. Predation, indexed by palatability, may select for increased flight speed and thoracic mass at the expense of the abdomen, but relative flight speed and thoracic mass were not significantly correlated.Within lineages (n=45 species for each sex), thoracic mass was uncorrelated with palatability in both sexes. Relative flight speed correlated positively with thoracic mass and negatively with body mass. Palatability and abdominal mass were negatively correlated for males but not females. Hence differences between the sexes in mass distribution suggest differences in reproductive constraints and predation stress.     

The impact of avian predator-scavengers on King Penguin Aptenodytes patagonicus chicks at Marion Island

While ashore King Penguins Aptenodytes patagonicus are fed upon by a guild of five predator-scavenger seabirds. During the winter (April-October) male Southern Giant Petrels Macronectes giganteus killed an estimated 6430 (11.2%) of Marion Island's King Penguin chicks, although most birds only scavenged in the colonies. The rate of predation varied, with peaks in April-May and in September. The proportion of successful attacks was 22.7%. There was a strong correlation between colony size and the rate of accumulation of chick corpses. Kelp Gulls Larus dominicanus and Lesser Sheathbills Chionis minor also scavenged penguin corpses but Northern Giant Petrels M. halli and female Southern Giant Petrels rarely entered the colonies. During the summer predation was mainly by Sub-Antarctic Skuas Catharacta lonnbergi which took eggs and small chicks.