Moulting patterns drive within-individual variations of stable isotopes and mercury in seabird body feathers: implications for monitoring of the marine environment

One major limitation in the use of body feathers of seabirds as a monitoring tool of the trophic structure and contamination levels of marine ecosystems is the degree of heterogeneity in feather chemical composition within individuals. Here, we tested the hypothesis that moulting patterns drive body feather heterogeneity, with synchronous moult minimizing within-individual variations, in contrast to asynchronous feather growth. Chicks of white-chinned petrels Procellaria aequinoctialis (representative of bird chicks) and adults of king penguins Aptenodytes patagonicus (representative of adult penguins) that moult their body feathers synchronously showed very low within-individual variations in their feather δ¹³C and δ¹⁵N values and mercury (Hg) concentrations. By contrast, body feathers of adults of Antarctic prions Pachyptila desolata (representative of adult seabirds with asynchronous feather growth during a protracted moult) presented much higher within-individual variances for the three parameters. These findings have three important implications for birds presenting a synchronous body moult. (1) They suggest that all body feathers from the same individual have identical δ¹³C and δ¹⁵N values and Hg content. (2) They predict negligible within-individual variations in the body feather values of other useful stable isotopes, such as δ²H and δ³⁴S, as well as in the concentrations of other compounds that are deposited in the keratin structure. (3) Analysis of one or any number of pooled body feathers is equally representative of the individual. In conclusion, we recommend that long-term routine monitoring investigations focus on birds presenting synchronous rather than asynchronous moult of body feathers both in marine and terrestrial environments. This means targeting chicks rather than adults and, for seabirds, penguins rather than adults of flying species.     

Interactions between sea urchins (Strongylocentrotus droebachiensis) and their predators in field and laboratory experiments

Field observations and manipulative experiments in a nearshore cobble bed (2 to 3 m below mean low water) at Eagle Head, Nova Scotia, Canada, between 1984 and 1986, showed that small juveniles ofStrongylocentrotus droebachiensis (3 to 6 mm diam) sheltering beneath cobbles had a refuge from predators such as rock crabs, small lobsters, and fish. Sea urchins gradually outgrew these refuges and small adults (25 to 30 mm) required larger rocks as shelter from predators, particularly large cancrid crabs. Small juveniles were usually solitary and well dispersed beneath cobbles, whereas small adults tended to aggregate on the undersides and in the interstices of boulders. These aggregations may develop passively as sea urchins accumulate in suitablysized refuges. Chemotaxis experiments indicate that juvenileS. droebachiensis are repelled by waterborne stimuli from conspecifics. In a factorial experiment, effects of the presence of potential predators (rock crabs and lobsters) and/or food (kelp) on the behaviour of large juvenile (10 to 15 mm) and small adult sea urchins were examined in flowing seawater tanks. Both size classes formed exposed feeding aggregations when kelp was provided as food, irrespective of the presence or absence of predators. In the absence of kelp, each size class responded differently to the presence of a predator: juveniles became more cryptic, whereas adults aggregated on the tank sides. Increased movement to the sides of a tank in the presence of a predator may reflect a flight response, since chemotaxis experiments indicated thatS. droebachiensis is repelled by waterborne chemical stimuli from predators. Observational and experimental data suggest that predation at the late juvenile and early adult stages may influence population structure, distribution and abundance ofS. droebachiensis.     

Costs of overlapping reproduction and moult in passerine birds: an experiment with the pied flycatcher

The process of moult in birds requires the investment of substantial amounts of energy and nutrients in feather production and attendant processes. Flight performance may be reduced during the moult of the wing feathers, and moulting birds may suffer from an increased predation risk. These factors may explain why the moult is usually separated in time from other energetically demanding processes, such as reproduction. In this study, we investigated the importance of the temporal separation of moult from breeding activities in the pied flycatcher Ficedula hypoleuca. We induced a moult-breeding overlap by removing the two innermost primaries on both wings, thus imitating the natural loss of these feathers during the initial stage of the moult. The experiment probably did not stimulate feather regrowth, but may have reduced flight performance. Just before fledging of the young, manipulated males and females, as well as the fledglings, had a lower body condition than control birds. The return rate of adult males in the year following the manipulation was significantly lower for males in the manipulated group than for control males. The return rate of females was lower than that of males, but there was no difference between the two female groups. The number of nestlings recruited in the year after the manipulation was significantly lower for the group with an induced moult-breeding overlap than for the control group. The reproductive performance in recruited and returning old females was independent of the manipulation, and in returning adults and recruits the feather quality in terms of number of barbs/mm was independent of the manipulation. These results suggest that a moult-breeding overlap induces fitness costs in terms of reduced recruitment rate and adult survival in a long-distance migrant. Received: 17 September 1997 / Accepted after revision: 23 February 1998     

Population dynamics of the commensal spider crab Inachus phalangium (Decapoda: Maiidae)

The ecology of the spider crab Inachus phalangium (Fabricius, 1775) (Decapoda: Maiidae) was studied in the field. I. phalangium inhabits the sublittoral on the sea anemone Anemonia sulcata Pennant. From July 1981 to April 1984 in the Mediterranean (Banyuls sur Mer, southern France) more than 3000 anemones were examined and ca 1800 I. phalangium were found on them. The population dynamics' generation cycles, reproductive activities and the dynamics of the sex ratio were investigated. The density of juveniles (crabs before the pubertal moult) on anemones changed in a yearly cycle from low in the first six months to very high in the second six months. The first occasional young crabs of a generation appeared in March/April (3rd and 4th decapodite stages) on the anemones. Their density increased enormously in the following months. The generation grew gradually on the anemones and moulted into puberty in September-January. Density of adults (crabs after the pubertal moult) on anemones changed in a yearly cycle from low to high from the summer to winter months. A new adult generation was recruited every autumn through the pubertal moult and disappeared in the following summer. Female reproductive activity continued throughout the year. Females carried several broods in succession, but the frequency of breeding females fluctuated on a yearly cycle. The highest percentage of egg-carrying females, i.e. the peak of the reproductive season, lay in the first half of the year. The maximal life span of a generation, from the hatching of the first larvae to the disappearance of the last adults, lasted 1.5 to 2 years. Males moulted into puberty ca one month later than females. The moulting distribution of adult males had roughly the same course as in females. An adult male generation diet out about one to two months before the female generation. Life expectancy was therefore 14 to 17 months for females and 12 to 15 months for males. The sex ratio of juveniles shortly before the pubertal moult was balanced. The sex ratio of adults shifted from 1:1 at the beginning of the reproductive period to ca 1:9 in favour of females at its end.     

The influence of kinship on foraging competition in Siberian jays

Foraging competition in Siberian jay groups was examined in relation to dominance and kinship to determine whether juvenile offspring, by associating with adults, gained in food acquisition relative to juvenile immigrants. Members of the adult pair were dominant over juvenile cohort members and males were dominant to females, although an inter-sexual hierarchy, with male juveniles occasionally overlapping adult females, was suggested. Few competitive asymmetries were found between adults and retained offspring or adults and immigrant juveniles when they were competing for food together, but in kin and non-kin foraging groups, respectively. Male offspring visited the bait site more frequently than adult males, and female immigrants spent less time at the bait site than adult females. Under these circumstances, hoarding activities may limit the ability of alpha members to control resources. In mixed groups containing both juvenile offspring and juvenile immigrants, no difference was found in the number of visits made to the bait site, although load sizes and foraging rates were lower for immigrant birds. Retained juveniles obtained greater load sizes and foraging rates when associating with adults. The social dominance of parents suggests that they control juvenile foraging. Although offspring benefit in the presence of adults, adults may incur a cost to their restraint by spending more time at the bait site when competing with immigrants. These results extend conclusions from previous work describing the role of selective tolerance by adults which relaxes competition with retained offspring in Siberian jay winter groups. The present findings suggest that offspring benefit in both immediate and future energy gains, which may have a direct influence on survival. Received: 18 September 1996 / Accepted after revision: 26 January 1997     

Dominance,prior occupancy,and winter residency in the great tit (Parus major)

Summary This study reports an aviary experiment aimed at determining what affects social dominance in the great tit (Parus major), especially why older birds (adults) in nature normally dominate younger ones (juveniles). When birds were matched with respect to age, prior residency determined dominance. Without a difference in prior residency older birds dominated younger ones. However, when juvenile birds had a prior residency advantage over adult birds, they often became dominant. This was especially so when the juvenile bird was large relative to the adult bird. When a resident juvenile male was also consorted by a female, he became dominant over an adult male on most occasions. An experiment where the dominant bird was removed and later returned to the aviary failed to produce more than one shift in dominance. However, the proportion of reversals in dominance interactions increased with separation time. It is argued that the fact that dominance depends on prior residency selects for winter residency in the great tit.Offprint requests to: H.G. Smith     

Responses to conspecific scent marks and the ontogeny of territorial marking in immature terrestrial salamanders

We investigated the ontogeny of responses to scent marks in immature terrestrial salamanders (Salamandra lanzai) that inhabit the southwest Alps. In this species, sexual maturity is usually reached at about 8 years, and adults exhibit territoriality. One should expect territoriality to take place largely before the acquisition of sexual maturity if sexual competition is not the main force driving territoriality. However, both the difficulties for inexperienced animals to find a suitable territory and size-related competition may delay the acquisition of territoriality in juveniles. We performed choice tests with juveniles belonging to two age groups (1±1 years old versus 4±1 years old). Each focal animal was offered, in random order, the choice between two shelters, one without scent and the other containing one of the following scents: own, same-aged animal, a juvenile belonging to the other age group, or adult female. We also performed choice tests with adult females for which the scents of two juveniles belonging to a different age group were successively offered in a random order. Older juveniles were strongly attracted toward their own shelter and mostly avoided the shelters that contained the scent of juveniles of about the same age and of adult females. Adult females avoided the scents of older juveniles but not younger juveniles. These results therefore suggest that older juveniles use territorial marking. Conversely, young juveniles behaved randomly with respect to their own scents and to those of a same-aged juvenile, and they were significantly attracted towards the odor of an older animal, especially adult females. Both older juveniles and adult females displayed a random behavior toward the scents of young juveniles. Our results suggest that young juveniles do not defend territories but use spaces occupied by older individuals.Communicated by A. Mathis     

Plasticity versus repeatability in seabird migratory behaviour

Pelagic seabird populations can use several discrete wintering areas, but it is unknown if individuals use the same wintering area year after year. This would have consequences for their population genetic structure and conservation. We here study the faithfulness of individuals to a moulting area within and among years in a small pelagic seabird, the Thin-billed prion, which moult their primary feathers during the early part of the non-breeding period. According to stable carbon isotope ratios (δ¹³C) of these feathers, 90% of Thin-billed prions moult in Antarctic and 10% in South American waters. Repeated samples from individuals in 2 or 3 years indicated that several birds changed between Antarctic and South American moulting areas or vice versa. However, individuals moulting in an area in one year were more likely to do so again. Four out of five adults maintained highly conserved δ¹³C over the extended moulting period. One bird, however, had systematic changes in δ¹³C indicating latitudinal movements between the two areas during moult. Thus, the present data show that this seabird species has a highly flexible migratory strategy, not only at the population level, but also at the individual level, enabling these seabirds to exploit a highly unpredictable environment.     

Endogenous control of timing of metamorphosis in megalopae of the shore crab Carcinus maenas

Batches of hundreds of freshly collected megalopae of the shore crab Carcinus maenas (L.) showed persistent circatidal rhythms of moulting to the juvenile crab stage when maintained in constant laboratory conditions. Peaks of moulting occurred around expected times of high tides, with few megalopae moulting at other times. In larvae collected offshore, the highest tidally-timed peak of metamorphosis occurred during the second to fifth expected times of high tide, and metamorphosis of 50% of each batch took about 22 h or longer. In contrast, in larvae collected at the water's edge, 70% metamorphosed during the first expected episode of high tide, within 6 to 8 h after collection. However, although inshore megalopae moulted before offshore ones, the tidal timing of moulting remained unaltered whether megalopae were collected at neaps or springs, from the water's edge or farther offshore, in the presence or absence of natural substratum, and under various light–dark and salinity regimes. Metamorphosis of C. maenas megalopae around the times of high tides may enhance settlement into the upper intertidal zone. Early juveniles of the crab apparently prefer that zone as they are most abundant there and, unlike adults, do not undertake up-and-down-shore migration with tides. The present finding demonstrates, for the first time, endogenous physiological timing of circatidal periodicity in the metamorphic moult of crab megalopae, suggesting that endogenous factors, as well as exogenous ones should be taken into account in considering the process of settlement by crab megalopae. Received: 21 February 1996 / Accepted: 27 November 1996     

Dominance behaviour,body weight and fat variations,and partial migration in European blackbirds Turdus merula

Summary A partially migratory population of European blackbirds Turdus merula was studied in central Europe during the non-breeding season. Body weight and the amount of subcutaneous fat deposits showed seasonal variations typical for each age and sex category. Juveniles were fattest in autumn and spring and both juveniles and females lost fat and body weight in mid-winter. Adults and males reached their maximum body weight in mid-winter. These findings together with data from Schwabl (1983) suggest that juveniles and females constitute the main fraction of those birds migrating from the area in autumn. In winter mainly adults and males have access to food resources. These patterns may be regulated by dominance behaviour, since adults and males are usually dominant over juveniles and females throughout the non-breeding season. Subordinate birds that stay in the area probably suffer from a higher winter mortality but have the advantage of early breeding.     

Differences in herbivore preferences, phlorotannin production, and nutritional quality between juvenile and adult tissues from marine brown algae

 Juvenile and adult marine organisms differ in their morphology, chemistry, physiology, behavior, and ecology. Because juvenile algae are thinner, smaller, and have more delicate tissues than adults, they are often assumed to be more susceptible to grazers. We examined within-species food preferences of four common generalist herbivores for juvenile and adult tissues of eight common brown algae in two-choice laboratory food-preference experiments. Our results showed that juvenile algae did not tend to be a preferred food of herbivores. Juvenile tissues were significantly preferred over adult tissues in only four of the 32 combinations of algae and herbivores tested. In 12 experiments, adult tissues were preferred over juvenile tissues, and no choice occurred in the remaining 16 experiments. When sea urchins exhibited a preference, it was always for adult tissues. The other three herbivores, an isopod and two snails, were more variable in their choices, sometimes preferring juveniles, sometimes adults, and sometimes having no preference. We measured nitrogen and phlorotannin concentrations in adult and juvenile seaweeds to see whether these parameters were correlated with herbivore food preferences. Nitrogen levels were similar in juveniles and adults of three algal species and were higher in juveniles of two. Phlorotannin levels were higher in juveniles of four species and lower in juveniles of one. The other three species showed no differences in phlorotannin levels. Phlorotannin concentrations decreased with increasing juvenile size in three species and increased with increasing size in one species. Neither nitrogen nor phlorotannin concentrations explained overall herbivore food preferences for algae of different stages. Our results suggest that preferences of certain grazers for juvenile algae are not as strong as previously assumed and are dependent on herbivore species. Preferences between juveniles and adults are likely to be determined by a combination of morphological and chemical features of the tissues and the unique responses of herbivore species to those features. Received: 10 April 2000 / Accepted: 19 November 2000     

Ontogeny of space use and diet of two temperate damselfish species, <Emphasis Type="Italic">Parma microlepis</Emphasis> and <Emphasis Type="Italic">Parma unifasciata</Emphasis>

Studies of reef fish herbivory have mainly focused on the impacts and behaviour of adults of tropical species. In this study, the ontogenetic shifts in home range, aggression, feeding rate, diet and gut morphology in juveniles and adults of two temperate territorial damselfishes, Parma microlepis and Parma unifasciata, were determined. Both P. microlepis and P. unifasciata juveniles under 80 mm TL exhibited no aggressive chases towards conspecifics or other species, while above 80 mm TL aggressive chase frequency increased in conjunction with an increase in home range, defended as a territory. Ontogenetic diet shifts, characterised by an increase in herbivory (P. unifasciata: juveniles: 64% plant material, adults: 95% plant material; P. microlepis: juveniles: 43% plant material, adults: 67% plant material) were observed for both species. The ratio of digestive tract length to body length, which often accompanies a switch to herbivory, increased significantly with ontogeny for both species. Compared to tropical confamilial grazers, these temperate damselfish species feeding rates were lower, and they had larger territories which were not as strongly defended (fewer aggressive chases).     

Dispersal, pair formation and social structure in gibbons (Hylobates lar)

We report observations on reproduction, natal dispersal, pair formation, and group structure based on longitudinal observations of several white-handed gibbon (Hylobates lar) groups spanning 18 years. Our observations are at odds with the traditional view that gibbons live in nuclear family groups consisting of a pair of adults and their offspring, and that parents exclude young from the family territory when they reach adult size. In the relatively dense Khao Yai study population, dispersing young usually obtain mates by replacing adults in existing territories, which creates non-nuclear families. Six subadults, five males and one female, matured and dispersed at an average age of 10 years, or about 2 years after reaching adult size. Average natal dispersal distance was 710 m, or between one and two territories away. At least two dispersing males replaced adults in neighboring groups. In one case, forcible displacement of the resident male resulted in a group which included a young juvenile presumably fathered by the previous male, two younger juveniles (probably brothers) from the new male's original group, and (later) offspring of the new pair. Social relations within this heterogeneous group remained harmonious: the adults groomed all the young and play occurred between all preadult members. In only two out of a total of seven cases of dispersal seen did two subadults pair and disperse into new territorial space. Nonreproducing subadults which delay dispersal may be tolerated by the adults provided that they contribute benefits to the adults or their offspring. Possible benefits include behaviors such as grooming, social play with juveniles, and support of the adult male in defending the territory. Delayed dispersal is probably advantageous in a saturated environment where there is no room for floaters, but subadults may also gain indirect fitness benefits by aiding siblings and other relatives. Received: 24 January 1997 / Accepted after revision: 12 January 1998     

Juvenile–adult relationship in the gregarious ophiuroid<Emphasis Type="Italic"> Ophiothrix fragilis</Emphasis> (Echinodermata): a behavioral and morphological study

Ophiothrix fragilis forms dense beds in the North Sea–English Channel region, where juveniles are exclusively found on adults. The aim of this study was to see how the behavior and morphology of juveniles could help elucidate the close juvenile–adult relationship found in this species. Juveniles are found on the disk, arms and in the bursae of adult conspecifics, the ones on the disks being significantly larger. No clear advantage seems to be gained by the juveniles being in the bursae, and their presence there is most likely due to juvenile movement on adults and to and from adults. Hooked spines serve as anchory organs during the early life of the juvenile, but growth of the arms enhances its anchory capabilities and the hooked spines become secondary in that respect. They regress as ophiuroids become older, to the advantage of the other spines used in suspension feeding. Juveniles are attracted to conspecifics, and true gregarious behavior has been observed. The tip of each arm, the terminal tentacle, plays a major role in distance chemoattraction, with juveniles needing at least one intact terminal tentacle to be able to initiate a response. Electron microscope observations of the terminal tentacle permitted us to recognize two different potential receptor structures designated stäbchens. The first possesses one long projecting cilia and is mostly present around the base of the terminal tentacle, while the other has one to five short projecting cilia and is mostly found on the tip. No receptors are found on the shaft. Receptors are not associated with secretory structures. Juveniles and adults are closely associated with one another, and both the morphology and behavior of juveniles play an important role in that relationship.Communicated by O. Kinne, Oldendorf/Luhe     

Winter migration of magellanic penguins (Spheniscus magellanicus) from the southernmost distributional range

A total of 22 magellanic penguins (Spheniscus magellanicus) from Isla Martillo in the Beagle Channel, Argentina, were successfully satellite tracked in 2004 (n = 7), 2005 (n = 7) and 2006 (n = 8) to monitor their winter migration after moult. Only one magellanic penguin migrated northwards into the Pacific Ocean, whereas all others remained in the Atlantic Ocean. In general, these birds left the island in an easterly direction, rounded Cabo San Diego, the southeasterly tip of South America, and continued northwards occupying inshore waters mostly less than 50 km from the coast, only occasionally venturing further offshore. By the end of the transmission period, birds were still travelling northwards and the most northerly positions were obtained from birds located in the area of Peninsula Valdés, Argentina, at a latitude of around 42°S, some 1,500 km from their breeding site on Isla Martillo. The mean maximum distance to the breeding site was, however, only 624 ± 460 km. The mean minimum distance covered during the study period was 1,440 ± 685 km, which corresponded to a mean distance of 23.2 ± 6.6 km covered per day. The northbound migration of the penguins could be separated into periods of rapid movement, interspersed with periods during which the birds remained for some time in particular coastal regions. Areas with a high density of daily penguin positions were observed in three distinct areas: at the northeastern coast of Tierra del Fuego, at the southern entrance of Golfo San Jorge and to the northeast of the Peninsula Valdez. The observed migration pattern is presumably driven by the formation and subsequent dispersal of areas of enhanced productivity as the season progressed. Our findings also suggest that magellanic penguins are increasingly threatened by human activities in coastal areas as penguins migrate northwards.     

Juvenile–adult associations in sea urchins ( Strongylocentrotus franciscanus and S. droebachiensis ): protection from predation and hydrodynamics in S. franciscanus

Juvenile red sea urchins, Strongylocentrotus franciscanus, aggregate under adult conspecifics, whereas sympatric juvenile green sea urchins, Strongylocentrotus droebachiensis, are typically more solitary and dispersed. Neither the potential advantage of juvenile sheltering nor the differences in post-settlement behavior between the two species has been demonstrated experimentally, but may be related to protection from predators and/or hydrodynamics. In predation experiments, juvenile vulnerability differed in the two species as the seastar Pycnopodia helianthoides consistently chose juvenile S. franciscanus over S. droebachiensis (100% vs. 0%). When associated with adults, juvenile mortality decreased dramatically in S. franciscanus (90% alone vs. 5% with adults), but very little in S. droebachiensis (85% vs. 75%). Not surprisingly, juvenile behavioral responses in the two species reflect this difference in vulnerability. Juvenile S. franciscanus sheltered under adults when predation risk was high, but not when risk was low (44% vs. 13%), whereas sheltering in S. droebachiensis was infrequent and not related to predation risk (7% for high risk versus 5% for low risk). From a hydrodynamic perspective, the presence of an adult led to the creation of a hydrodynamic refuge for juveniles, where average water velocities were reduced by > 60% around the adult urchin. Again, striking differences in sheltering rate were apparent in S. franciscanus (52% vs. 13% for high flow and low flow, respectively), but not S. droebachiensis (5% for high flow versus 4% for low flow). Sheltering behavior was also species-specific as juveniles did not shelter at high rates under adults of the opposite species (≤ 16%). A field survey confirmed these finding in that juvenile S. franciscanus abundance was associated with both adults and water motion (R ² = 0.80, P = 0.008, best-subsets regression). These results suggest that sheltering confers juvenile S. franciscanus with a degree of protection from predators and water motion, and that species-specific differences in this post-settlement behavior may be related to the differences in the protection afforded by adults.     

Use of Stress‐Hormone Levels and Habitat Selection to Assess Functional Connectivity of a Landscape for an Amphibian

Abstract: The influence of landscape matrix on functional connectivity has been clearly established. Now methods to assess the effects of different land uses on species’ movements are needed because current methods are often biased. The use of physiological parameters as indicators of the level of resistance to animal movement associated with different land uses (i.e., matrix resistance) could provide estimates of energetic costs and risks to animals migrating through the matrix. To assess whether corticosterone levels indicate matrix resistance, we conducted experiments on substrate choice and measured levels of corticosterone before and after exposure of toads (Bufo bufo) to 3 common substrates (ploughed soil, meadow, and forest litter). We expected matrix resistance and hormone levels to increase from forest litter (habitat of the toad) to meadows to ploughed soil. Adult toads had higher corticosterone levels on ploughed soil than on forest litter or meadow substrates. Hormone levels did not differ between forest litter and meadow. Toads avoided moving onto ploughed soil. Corticosterone levels in juvenile toads were not related to substrate type; however, hormone levels decreased as humidity increased. Juveniles, unlike adults, did not avoid moving over ploughed soil. The difference in responses between adult and juvenile toads may have been due to differences in experimental design (for juveniles, entire body used to measure corticosterone concentration; for adults, saliva alone); differences in the scale of sensory perception of the substrate (juveniles are much smaller than adults); or differences in cognitive processes between adult and juvenile toads. Adults probably had experience with different substrate types, whereas juveniles first emerging from the water probably did not. As a consequence, arable lands could act as ecological traps for juvenile toads.     

Mortality-minimizing sandpipers vary stopover behavior dependent on age and geographic proximity to migrating predators

Ecological theory for long-distance avian migration considers time-, energy-, and mortality-minimizing tactics, but predictions about the latter have proven elusive. Migrants must make behavioral decisions that can favor either migratory speed or safety from predators, but often not both. We compare the behavior of adult and juvenile western sandpipers Calidris mauri during the course of their temporally segregated passages at a major stopover site. Here, the passage and winter arrival of an important predator, the peregrine falcon Falco peregrinus begins near the end of the adult sandpiper passage (July) and increases rapidly through the juvenile passage (August). The mortality-minimizing hypothesis predicts that as the falcon front is distant but approaching, sandpipers should initially increase the fuel-loading rate (lowered vigilance and predator apprehension) to increase migration speed and so maintain their head start. As the falcon front gains proximity to and passes over the stopover site, sandpipers should become increasingly cautious. Our measurements show that adults decreased vigilance during the period prior to falcon arrival, and had lower vigilance overall than juveniles. Juveniles were more apprehensive, flying further and longer in response to disturbance by a falcon silhouette. This trend was reversed in response to a human approach. Both groups were more vigilant and more apprehensive in a study year with earlier falcon arrival. These results suggest that late (juvenile) and early (adult) migrants minimize mortality on migration in different ways, adults by increased migratory speed at the expense of caution on stopover sites, and juveniles by increased caution at the expense of speed.     

Characterisation of cysteine proteinase activities in the digestive tract of juvenile <Emphasis Type="Italic">Paragnathia formica</Emphasis> isopods,ectoparasites of estuarine fish

Juveniles of Paragnathia formica Hesse (Isopoda; Gnathiidae) are haematophagous ectoparasites, feeding on fish blood which supplies the nutrients for their development through growth and moulting, and the subsequent survival and reproduction of the free-living adults. Little is known of the mechanisms of digestion in juvenile gnathiids, and biochemical studies on the digestive tract of these interesting estuarine isopods have not been undertaken previously. Here, functionally active cathepsin-like cysteine proteinases are identified in the digestive system of juvenile praniza (fed) and zuphea (unfed) forms. The physiological pH of the digestive tract and the optimum proteolytic activities detected in praniza 3 homogenates using the cathepsin B/L, cathepsin B, and cathepsin H fluorogenic substrates, N-Carbobenzoxy-Phe-Arg-4-methoxy-2-naphthylamine (Z-phe-arg-MNA), N-Carbobenzoxy-Arg-Arg-4-methoxy-2-naphthylamine (Z-arg-arg-MNA) and Arg-4-methoxy-2-naphthylamine (H-arg-MNA), respectively, are in the acidic range (pH 5.8–6.7). Inhibition profiles against Z-phe-arg-MNA and Z-arg-arg-MNA using the cathepsin B inactivator urea, and cysteine proteinase inhibitors, support the presence of cathepsin L- and B-like enzymes. These proteolytic activities are 10–50 times higher in homogenates of praniza 3 compared with zuphea 3 forms. Histochemistry of praniza 3 sections reveals that the predominant enzyme activity towards Z-phe-arg-MNA is limited to the digestive glands during early and mid stages of digestion; later, this activity appears in the lining of the anterior hindgut. Moreover, activity towards Z-arg-arg-MNA is generally restricted to the digestive glands, and only occasionally present in the anterior hindgut. In conclusion, the digestive glands are the main site of cathepsin-like cysteine proteinase activities in P. formica juveniles; these enzymes appear to be important to the digestion of host fish blood enabling development through to the free-living adult.     

Cooperative breeding in purple gallinules: the role of helpers in feeding chicks

Summary The participation of breeders and helpers in the feeding of 21 broods of chicks was studied in a population of cooperatively breeding purple gallinules (Porphyrula martinica). In the breeding group, all birds over the age of 2 months fed chicks. Female breeders fed chicks at the highest rate, followed by male breeders and adult helpers, old juvenile helpers, and young juvenile helpers. The amount of food breeders fed chicks was independent of the number of helpers in the breeding group. However, breeders made fewer feeding visits when they had helpers. Male and female breeders spent similar amounts of time feeding chicks. Helpers spent significantly less time feeding chicks than did breeders. As helpers grew older they fed chicks at a faster rate, made more feeding visits and spent more time feeding chicks. Analysis of variance was used to determine which variables explained the variation in the brood feeding rate (amount of food delivered to an entire brood during one observation). Age of chicks had a significant nonlinear effect, and size of brood and number of helpers had significant linear effects on the brood feeding rate. Chicks in groups with helpers received more food and were accompanied for longer periods of time than chicks in groups without helpers; either or both of these factors may have led to increased chick survival.