Effect of sex ratio on growth and survival in Octopus vulgaris (Cuvier, 1797) reared in floating cages

[EN]Octopus vulgaris is a potential candidate to diversify European aquaculture for its rapid growth and high market prices (Vaz Pires et al. 2004). One factor affecting industrial development of octopus culture is sexual maturation under rearing conditions. Octopus females can lose up to 30-60% of their initial body weight during egg-laying (Iglesias et al., 2000) and die after the paralarvae hatch (Guerra,1992), while a correlation between males death and spermatic sac depletion has being recently reported by Estefanell et al. (2010b). The present experiment discusses the effect of three different sex ratios on growth, sexual maturation and survival in O. Vulgaris. Conclusions: Discarded bogue from fish farms could be used as alternative diet for the final stage of O. vulgaris ongrowing ; Male segregation would maximize biomass increment ; Under the conditions described, sex ratios close to 1:1 produced higher biomass increment than 4:1

Nuove tecniche per l'allevamento massale di Ditteri del genere Coenosia Meigen

The species of the genus Coenosia Meigen are polyphagous predators both in the larval and in the adult stage. In Europe five Coenosia species of the ‘tigrina group’ are naturally present in greenhouses, where they can establish for long periods. As their wide preys range includes important pests of protected crops, as Aleyrodidae, Sciaridae and Agromyzidae, Coenosia species are considered promising potential biological control agents. A method for rearing Coenosia species in vivo was developed for the first time in 1993 in Germany, where C. attenuata, C. strigipes and C. humilis were bred on Bradysia paupera (Diptera Sciaridae), reared on Fusarium spp. cultivated on wood fibre. Although this method was partially simplified afterwards, it is still too complex and expensive for a mass production. This research aimed at simplifying this rearing procedure and making it cheaper, in the perspective of an eventual mass production. Studies on potential preys were conducted, to determine their suitability for C. attenuata larvae and adults and to develop rearing methods. Biology and rearing methods of Bradysia paupera Tuomikoski, Scatella stagnalis Fallén and Drosophila melanogaster Meigen (Diptera: Sciaridae, Ephydridae, Drosophilidae) were compared. B. paupera resulted the most suitable prey for rearing C. attenuata in vivo. The Sciarid fly was effectively reared on damp coconut fibre with fresh Agaricus bisporus (J.E. Lange) Pilát, thus simplifying the existing method. After preliminary trials with different potential preys, attempts to rear C. attenuata in vivo on B. paupera and D. melanogaster were made. The best results were obtained with B. paupera, reared on coconut fibre and A. bisporus, but the method needs further improvement. Trials of in vitro rearing of C. attenuata were also made: as no specific diet for Coenosia species is reported in literature, different potentially suitable media were tested. Among these, a specific diet for Diptera Tachinidae resulted a good starting point for further studies and improvements. The biology of C. attenuata adults captured in greenhouses was also studied, by observing both groups and isolated individuals. Data on lifespan, daily number of preys per adult, daily number of laid eggs and hatching rate were recorded, and the effects of different foods on these parameters were analyzed. The following foods were compared: D. melanogaster adults only, as preys for C. attenuata; D. melanogaster adults and a water-honey solution; the water-honey solution only. Honey resulted an effective food integration for C. attenuata, increasing lifespan and the number of egg laying females. It is possible that in greenhouses Coenosia adults complete the preys diet with nectar and/or honeydew. Moreover, the integration with honey reduced the daily preys consumption. This may allow to prevent cannibalism among Coenosia adults in the rearing conditions, where high population densities are required. A survey of the Coenosia species naturally present in Lombardy greenhouses was conducted. The species C. attenuata, C. strigipes, C. tigrina and C. atra were detected. C. attenuata resulted the most common, recorded in most greenhouses and for consecutive years. Besides, the presence of potential preys, weeds and the crops were recorded in each greenhouse. Nevertheless, it is difficult to determine the relation between these parameters and the presence of Coenosia species.

Die Prozessierung des humanen Amyloid-Vorläuferproteins im Verlauf der replikativen Seneszenz

Die Alterung stellt den größten Risikofaktor für die Entwicklung der Alzheimer Krankheit dar, wobei die biochemische Basis dieser Korrelation bisher nicht bekannt ist. Ein möglicherweise zentraler Mechanismus der Alzheimer Pathologie wird durch die Prozessierung von APP repräsentiert, die in der Synthese von Aβ resultiert. Der Einfluss zellulärer Alterung auf die Biochemie der APP-Prozessierung ist bislang weitestgehend ungeklärt. In der vorliegenden Arbeit wurde gezeigt, dass die Prozessierung von endogenem APP im Verlauf der Zellalterung humaner Fibroblasten progressiv verringert wird. Die Bildung der intrazellulären APP-Spaltfragmente (C99, C83 und AICD) nahm mit zunehmender Lebensspanne ab und war gleichfalls mit einer reduzierten Synthese von extrazellulären APP-Fragmenten (sAPP, sAPPα) verbunden. Weiterhin wurde nachgewiesen, dass die Reifung von APP in seneszenten Zellen selektiv reduziert war, und dass dies durch altersabhängig erhöhte zelluläre Cholesterolspiegel vermittelt wurde. Von den APP-prozessierenden Sekretasen waren die Proteinspiegel von Presenilin-1 und Nicastrin, beides Komponenten der γ-Sekretase, im Verlauf der Zellalterung graduell verringert. Dies hatte einen progressiven Rückgang der enzymatischen Aktivität der γ-Sekretase zur Folge, wodurch die Prozessierung von APP unmittelbar reduziert wurde. Die Proteinspiegel von ADAM10, einer α-Sekretase, sowie der β-Sekretase, BACE, wiesen keine Altersregulation auf, aber interessanterweise wurde eine erhöhte enzymatische Aktivität der β-Sekretase in seneszenten Zellen nachgewiesen. Die γ-Sekretase sowie BACE sind in Lipid Rafts lokalisiert, geordneten Membransubdomänen, die hohe Cholesterol- und Caveolin-1-Spiegel aufweisen. Obwohl das Gesamtniveau dieser strukturellen Komponenten von Lipid Rafts in seneszenten Zellen erhöht war, war die Assoziation beider Moleküle mit Lipid Rafts reduziert und sie akkumulierten in speziellen Organellen, die höchstwahrscheinlich Lipidkörper darstellen. Somit wurde gezeigt, dass Lipid Rafts im Zuge der Zellalterung disintegrieren beziehungsweise in ihrem Gesamtspiegel reduziert waren. Diese altersabhängige Membranmodifikation war mit einer veränderten Verteilung von Presenilin-1 und BACE zwischen der Lipid Raft und der Nicht Raft Fraktion der Membran verbunden, die möglicherweise das Potential dieser Enzyme zur Prozessierung von APP reduzierte. In einem zweiten Teil der Arbeit wurden transgene C. elegans konstruiert, die humanes APP exprimieren, das C-terminal an GFP gekoppelt war. Diese Würmer wiesen eine reduzierte Fertilität, Eilegedefekte und eine verzögerte post-embryonale Entwicklung auf, die möglicherweise auf eine Transgen-vermittelte Neurodegeneration zurückgeführt werden können. Durch erste Untersuchungen der Prozessierung des Transgens konnten Spaltfragmente nachgewiesen werden, die potentiell auf eine spezifische Spaltung von APP durch die endogenen Sekretasen schließen lassen. Somit werden die Prozessierung sowie die Reifung von APP durch die altersabhängige Modifikationen zellulärer Biochemie nachhaltig beeinflusst. Zukünftige Studien sollen zeigen, ob sich diese zellulären Zusammenhänge in den Gesamtorganismus C. elegans übertragen lassen. Des Weiteren sollen die altersabhängigen zellulären Veränderungen, insbesondere des Cholesterol-Metabolismus und der Sekretaseaktivitäten, weitergehend analysiert werden, um zusätzliche Erkenntnisse über altersassoziierte Regulationen möglicher therapeutischer Ziele der Alzheimer Erkrankung zu gewinnen.

Managing invasive populations of Anoplophora chinensis and A. glabripennis in Lombardy

Two Asian longhorned beetles (Coleoptera: Cerambycidae), commonly known as Citrus Longhorned Beetle (CLB), Anoplophora chinensis (Forster), and Asian Longhorned Beetle (ALB), A. glabripennis (Motschulsky), are considered the most destructive wood borers introduced in Lombardy (northern Italy). This research aimed at (1) improving laboratory rearing methods for the biological control agent Aprostocetus anoplophorae (Hym.: Eulophidae), an egg parasitoid specific to CLB, and defining release techniques allowing its establishment; (2) test the efficacy of the sentinel tree technique for the early detection of CLB; and (3) evaluating the efficacy of traps baited with artificial lures in attracting adults of ALB and possibly CLB. Several problems were faced while rearing the egg parasitoid in laboratory. It appeared that the rate of parasitism of the hosts could depend on the age of the host eggs and/or age of the laying parasitoid females. Data results from the field experiments about A. anoplophorae release-capture showed that the percentage of slits containing a CLB egg was particularly low on most sentinel trees and the percentage of CLB eggs that were killed, because of natural predators, was high. Only one egg amongst those exposed was attacked by the released parasitoid. These negative results were anyway very useful, since they provided evidence and information on the type of host plants to be used, the time necessary for the exposure of the plants to the egg-laying CLB females, the number of laying parasitoid females to be inserted per cage. The sentinel trees technique revealed to be not successful; signs and symptoms of CLB presence were not recorded during the two seasons of field observations (2012-2013). Extremely positive was instead the trial with artificial lures carried out during summer 2013. A total of 32 beetles were captured (4 ALB and 28 CLB) deploying 50 baited traps.

Spawning coordination of mates in a shell brooding cichlid

Aim. External fertilisation requires synchronisation of gamete release between the two sexes. Adequate synchronisation is essential in aquatic media because sperm is very short-lived in water. In the cichlid Lamprologus callipterus, fertilisation of the eggs takes place inside an empty snail shell, where females stay inside the shell and males have to ejaculate into the shell opening. This spawning pattern makes the coordination of gamete release difficult. Methods. This study examined the synchronisation of males and females during egg laying. Results. The results showed that the male initiates each spawning sequence and that sperm release and egg laying are very well synchronised. 68% of all sperm releases occurred at exactly the same time when the female laid an egg, and 99% of ejaculations occurred within ±5 seconds from egg deposition. On average 95 eggs are laid one by one with intervals of several minutes between subsequent eggs, leading to a total spawning duration in excess of six hours. Conclusions. We discuss this exceptional spawning pattern and how it might reflect a conflict between the sexes, with males attempting to induce egg laying and females extending the egg laying period to raise the chance for parasitic males to participate in spawning.

Benefits of Induced Host Responses against an Ectoparasite

As a consequence of the deleterious effects of parasites on host fitness, hosts have evolved responses to minimize the negative impact of parasite infection. Facultative parasite-induced responses are favoured when the risk of infection is unpredictable and host responses are costly. In vertebrates, induced responses are generally viewed as being adaptive, although evidence for fitness benefits arising from these responses in natural host populations is lacking. Here we provide experimental evidence for direct reproductive benefits in flea-infested great tit nests arising from exposure during egg production to fleas. In the experiment we exposed a group of birds to fleas during egg laying (the exposed group), thereby allowing for induced responses, and kept another group free of parasites (the unexposed group) over the same time period. At the start of incubation, we killed the parasites in both groups and all nests were reinfested with fleas. If induced responses occur and are adaptive, we expect that birds of the exposed group mount earlier responses and achieve higher current reproductive success than birds in the unexposed group. In agreement with this prediction, our results show that birds with nests infested during egg-laying have (i) fewer breeding failures and raise a higher proportion of hatchlings to hedging age; () offspring that reach greater body mass, grow longer feathers, and hedge earlier, and (iii) a higher number of recruits and first-year grandchildren than unexposed birds. Flea reproduction and survival did not differ significantly between the two treatments. These results provide the first evidence for the occurrence and the adaptiveness of induced responses against a common ectoparasite in a wild population of vertebrates. [References: 50]

Cellular immune response, stress resistance and competitiveness in nestling great tits in relation to maternally transmitted carotenoids

1. Egg yolks contain carotenoids that protect biological molecules against free-radical damage and promote maturation of the immune system. Availability of carotenoids to birds is often limited. Trade-offs can thus arise in the allocation of carotenoids to different physiological functions, and mothers may influence the immunocompetence of nestlings by modulating the transfer of carotenoid to the yolk.;2. In the great tit Parus major, we experimentally manipulated the dietary supply of carotenoid to mothers, and partially cross-fostered hatchlings to investigate the effect of an increased availability of carotenoids during egg laying on immunocompetence of nestlings.;3. In addition, we infested half of the nests with hen fleas Ceratophyllus gallinae to investigate the relationship between carotenoid availability, resistance to ectoparasites and immunocompetence.;4. We found that the procedure of cross-fostering can reduce the immune response of nestlings, but this effect can be compensated by the maternally transferred carotenoids. Cross-fostered nestlings of carotenoid-supplemented females show a similar immune response to non-cross-fostered nestlings, while cross-fostered nestlings of control females mounted a weaker cell-mediated immune response. This suggests that yolk carotenoids may help nestlings to cope with stress, for example the one generated by cross-fostering and/or they may enhance nestling competitiveness.;5. There was no statistically significant interaction between parasite and carotenoid treatments, as would be expected if carotenoids helped nestlings to fight parasites. Under parasite pressure, however, lighter nestlings raised a lower immune response, while the immune response was only weakly correlated with body mass in uninfested nests.

Modulatory effects of bag cell peptides in {\it Aplysia}: Behavioral and electrophysiological studies

Reproductive hormones have effects on the nervous system not directly related to reproductive function. In the rat, for example, luteinizing hormone releasing hormone has dramatic effects on learning and memory. The present work attempts to examine the effects of reproductive hormones on non-reproductive behaviors and the neural loci and mechanisms underlying these effects in Aplysia, an animal whose behaviors, reproductive hormones and neural circuitry have been well characterized.^ In Aplysia, the neurosecretory bag cells release several peptides that are responsible for eliciting egg laying. The effects of these peptides on the defensive tail-siphon withdrawal reflex, as well as sensitization of this reflex, were examined. Sensitization, a simple form of nonassociative learning, refers to the behavioral enhancement of a response to a test stimulus after the presentation of a strong stimulus, that may last minutes (short-term) or days (long-term). An extract of the bag cells (BCE) inhibited the baseline siphon component of the tail-siphon withdrawal reflex and suppressed long-term, but not short-term, sensitization of the reflex. Preliminary experiments suggest that BCE also affects the tail component of the tail-siphon withdrawal reflex.^ To determine the neural mechanisms underlying the inhibition of the baseline reflex, electrophysiological studies were performed using an in vitro analogue of the tail-siphon withdrawal reflex to examine the ability of BCE, as well as the individual bag cell peptides (BCPs), to modulate the circuitry of the reflex. Bag cell extract attenuated the synaptic strength of the monosynaptic connections between tail sensory neurons and tail motor neurons. When individually applied only $\beta$-BCP produced a similar attenuation. This effect of $\beta$-BCP was not dependent on changes in duration of the presynaptic action potential.^ An in vitro analogue of long-term sensitization training was developed to examine the mechanisms by which the BCPs may affect long-term sensitization of the tail-siphon withdrawal reflex. This analogue exhibited both short- and long-term facilitation of the connections between the tail sensory and motor neurons.^ The results of these behavioral and electrophysiological experiments suggest that the BCPs inhibit the tail-siphon withdrawal reflex, at least in part, by modulating the synaptic strength of the connections between the sensory neurons and motor neurons underlying the reflex. One candidate for this effect is $\beta$-BCP. Thus, the peptides which elicit egg laying may also serve other functions such as the inhibition of defensive reflexes. In addition, these experiments raise the possibility that BCPs may exert a long lasting effect ($>$24 hr), suppressing long-term sensitization of the tail-siphon withdrawal reflex. ^

Man-induced activities modify demographic parameters in a long-lived species: effects of poisoning and health policies

Recent changes in sanitary policies within the European Union (EU) concerning disposal of carcasses of domestic animals and the increase of non-natural mortality factors, such as illegal poisoning, are threatening European vultures. However, the effects of anthropogenic activities on demographic parameters are poorly studied. Using a long-term study (1994–2011) of the threatened Pyrenean Bearded Vulture Gypaetus barbatus population, we assess the variation in the proportion of breeding pairs, egg-laying dates, clutch size, breeding success, and survival following a sharp reduction in food availability in 2005 due to the application of restrictive sanitary policies decreasing livestock carcass availability. We found a delay in laying dates and a regressive trend in clutch size, breeding success, and survival following policy change. The maintenance of specific supplementary feeding stations for Bearded Vultures probably reduced the negative effects of illegal poisoning and food shortages, which mainly affected subadult survival. A drop in food availability may have produced changes in demographic parameters and an increase in mortality due to an increased exposure to contaminated food. As a result, supplementary feeding as a precautionary measure can be a useful tool to reduce illegal poisoning and declines in demographic parameters until previous food availability scenarios are achieved. This study shows how anthropogenic activities through human health regulations that affect habitat quality can suddenly modify demographic parameters in long-lived species, including those, such as survival, with high sensitivity to population growth rate.

(Table 1) Stage of the lemming cycle and nesting parameters of the greater snow geese between 1996-2005, Bylot Island

Resource pulses are common in various ecosystems and often have large impacts on ecosystem functioning. Many animals hoard food during resource pulses, yet how this behaviour affects pulse diffusion through trophic levels is poorly known because of a lack of individual-based studies. Our objective was to examine how the hoarding behaviour of arctic foxes (Alopex lagopus) preying on a seasonal pulsed resource (goose eggs) was affected by annual and seasonal changes in resource availability. We monitored foraging behaviour of foxes in a greater snow goose (Chen caerulescens atlanticus) colony during 8 nesting seasons that covered 2 lemming cycles. The number of goose eggs taken and cached per hour by foxes declined 6-fold from laying to hatching, while the proportion of eggs cached remained constant. In contrast, the proportion of eggs cached by foxes fluctuated in response to the annual lemming cycle independently of the seasonal pulse of goose eggs. Foxes cached the majority of eggs taken (> 90%) when lemming abundance was high or moderate but only 40% during the low phase of the cycle. This likely occurred because foxes consumed a greater proportion of goose eggs to fulfill their energy requirement at low lemming abundance. Our study clearly illustrates a behavioural mechanism that extends the energetic benefits of a resource pulse. The hoarding behaviour of the main predator enhances the allochthonous nutrients input brought by migrating birds from the south into the arctic terrestrial ecosystem. This could increase average predator density and promote indirect interactions among prey.

Investigation of knot nests on Ellesmere Island

Knots arrive on Ellesmere Island in late May or early June. At Hazen Camp small flocks were present on 3 June 1966, but the main influx occurred 5 June when many flocks were seen ranging in size from 6 to 60 individuals. The sexes appeared to arrive together, but the manner of pair-formation was not determined. By 7 June pairs were distributed over the tundra with large feeding flocks forming at snowfree wet marshy areas. Most nests were on Dryas-hummocked slopes and tundra, either dry or moist, with some on clay plains and summits in a mixed Dryas and Salix vegetation. A census area of 240 ha supported at least 3 breeding pairs, and possibly 5; the total number of pairs breeding in the Hazen Camp study area was estimated to be about 25 (1.09 pairs/km**2). Egg-laying (4 nests) extended from 15 to 28 June, with 3 of the 4 sets completed between 20 and 23 June. Both sexes incubated, one of the pair more regularly than the other. The song-flight display of the male was performed most frequently during egglaying and incubation. The incubation period of the last egg in one clutch was established as being between 21.5 and 22.4 days. Four nests hatched between 12 and 20 July, and the hatching period of the entire clutch was less than 24 hours. Four of 7 nests (57 %) survived and egg survival (53 %) was low. Families left the nesting area so on after hatching, concentrating at ponds where food was readily available for the young. Both adults attended the young during the pre-fledging period, but the females apparently departed before the young had hedged. Males left once the young could fly and the adult fall migration was complete by early August. Most 01 the young departed belore mid-August. Fall migration is complete by late August or early September. The breeding season appears to be timed to peak load supply for the young. Adult Chironomidae emergence was highest between 3 and 17 July, the period during which most successful nests hatched. The increasing scarcity of adult insects for the young after mid-July was offset by family movements over the tundra and the early departure of half the adult population. Food also seemed to influence the distribution of breeding pairs aver the tundra, restricting them to the general vicinity of marshes, streams, and ponds where food is most available when the young hatch. Territoriality in the Knot appears to be closely associated with the protection of the nest against predators and has at least a local effect in regulating the number of breeding pairs. Plant material was important in the diet of adult Knots throughout the summer and the primary food from the time of arrival until mid-June. After mid-June the percentage of animal matter increased as dipterous insects became available (especially adult Chironomidae), but plant materials continued to constitute a large part of the diet, usually more than 50 %. The food of the young before fledging consisted principally of adult chironomids.

(Table 1) Reproductive performance of brown skua pairs with and without feeding territories at Potter Peninsula/King George Island from 1998/1999 to 2000/2001

In the maritime Antarctic, brown skuas (Catharacta antarctica lonnbergi) show two foraging strategies: some pairs occupy feeding territories in penguin colonies, while others can only feed in unoccupied areas of a penguin colony without defending a feeding territory. One-third of the studied breeding skua population in the South Shetlands occupied territories of varying size (48 to >3,000 penguin nests) and monopolised 93% of all penguin nests in sub-colonies. Skuas without feeding territories foraged in only 7% of penguin sub-colonies and in part of the main colony. Females owning feeding territories were larger in body size than females without feeding territories; no differences in size were found in males. Territory holders permanently controlled their resources but defence power diminished towards the end of the reproductive season. Territory ownership guaranteed sufficient food supply and led to a 5.5 days earlier egg-laying and chick-hatching. Short distances between nest and foraging site allowed territorial pairs a higher nest-attendance rate such that their chicks survived better (71%) than chicks from skua pairs without feeding territories (45%). Due to lower hatching success in territorial pairs, no difference in breeding success of pairs with and without feeding territories was found in 3 years. We conclude that skuas owning feeding territories in penguin colonies benefit from the predictable and stable food resource by an earlier termination of the annual breeding cycle and higher offspring survivorship.

At surface behaviour of emperor penguins from Pointe Géologie, Adélie Land, Antarctica, from expedition DDU 2005

The foraging distributions of 20 breeding emperor penguins were investigated at Pointe Géologie, Terre Adélie, Antarctica by using satellite telemetry in 2005 and 2006 during early and late winter, as well as during late spring and summer, corresponding to incubation, early chick-brooding, late chick-rearing and the adult pre-moult period, respectively. Dive depth records of three post-egg-laying females, two post-incubating males and four late chick-rearing adults were examined, as well as the horizontal space use by these birds. Foraging ranges of chick-provisioning penguins extended over the Antarctic shelf and were constricted by winter pack-ice. During spring ice break-up, the foraging ranges rarely exceeded the shelf slope, although seawater access was apparently almost unlimited. Winter females appeared constrained in their access to open water but used fissures in the sea ice and expanded their prey search effort by expanding the horizontal search component underwater. Birds in spring however, showed higher area-restricted-search than did birds in winter. Despite different seasonal foraging strategies, chick-rearing penguins exploited similar areas as indicated by both a high 'Area-Restricted-Search Index' and high 'Catch Per Unit Effort'. During pre-moult trips, emperor penguins ranged much farther offshore than breeding birds, which argues for particularly profitable oceanic feeding areas which can be exploited when the time constraints imposed by having to return to a central place to provision the chick no longer apply.

(Table 1) Breeding parameters for 60 male Adélie penguins (Pygoscelis adeliae) at station Dumont d'Urville (Antarctica) depending on treatment (control vs. CORT)

Corticosterone, the main stress hormone in birds, mediates resource allocation, allowing animals to adjust their physiology and behaviour to changes in the environment. Incubation is a time and energy-consuming phase of the avian reproductive cycle. It may be terminated prematurely, when the parents' energy stores are depleted or when environmental conditions are severe. In this study, the effects of experimentally elevated baseline corticosterone levels on the parental investment of incubating male Adelie penguins were investigated. Incubation duration and reproductive success of 60 penguins were recorded. The clutches of some birds were replaced by dummy eggs, which recorded egg temperatures and rotation rates, enabling a detailed investigation of incubation behaviour. Corticosterone levels of treated birds were 2.4-fold higher than those of controls 18 days post treatment. Exogenous corticosterone triggered nest desertion in 61% of the treated birds; consequently reducing reproductive success, indicating that corticosterone can reduce or disrupt parental investment. Regarding egg temperatures, hypothermic events became more frequent and more pronounced in treated birds, before these birds eventually abandoned their nest. The treatment also significantly decreased incubation temperatures by 1.3 °C and lengthened the incubation period by 2.1 days. However, the number of chicks at hatching was similar among successful nests, regardless of treatment. Weather conditions appeared to be particularly important in determining the extent to which corticosterone levels affected the behaviour of penguins, as treated penguins were more sensitive to severe weather conditions. This underlines the importance of considering the interactions of organisms with their environment in studies of animal behaviour and ecophysiology.

(Table 1) Nesting characteristics of greater snow geese (Chen caerulescens atlanticus) on Bylot Island between 1995-2005

1. Habitat heterogeneity and predator behaviour can strongly affect predator-prey interactions but these factors are rarely considered simultaneously, especially when systems encompass multiple predators and prey. 2. In the Arctic, greater snow geese Anser caerulescens atlanticus L. nest in two structurally different habitats: wetlands that form intricate networks of water channels, and mesic tundra where such obstacles are absent. In this heterogeneous environment, goose eggs are exposed to two types of predators: the arctic fox Vulpes lagopus L. and a diversity of avian predators. We hypothesized that, contrary to birds, the hunting ability of foxes would be impaired by the structurally complex wetland habitat, resulting in a lower predation risk for goose eggs. 3. In addition, lemmings, the main prey of foxes, show strong population cycles. We thus further examined how their fluctuations influenced the interaction between habitat heterogeneity and fox predation on goose eggs. 4. An experimental approach with artificial nests suggested that foxes were faster than avian predators to find unattended goose nests in mesic tundra whereas the reverse was true in wetlands. Foxes spent 3-5 times more time between consecutive attacks on real goose nests in wetlands than in mesic tundra. Their attacks on goose nests were also half as successful in wetlands than in mesic tundra whereas no difference was found for avian predators. 5. Nesting success in wetlands (65%) was higher than in mesic tundra (56%) but the difference between habitats increased during lemming crashes (15%) compared to other phases of the cycle (5%). Nests located at the edge of wetland patches were also less successful than central ones, suggesting a gradient in accessibility of goose nests in wetlands for foxes. 6. Our study shows that the structural complexity of wetlands decreases predation risk from foxes but not avian predators in arctic-nesting birds. Our results also demonstrate that cyclic lemming populations indirectly alter the spatial distribution of productive nests due to a complex interaction between habitat structure, prey-switching and foraging success of foxes.