Predation by Australian magpies (Gymnorhina tibicen) on pasture invertebrates: Are non-territorial birds less successful?

Diet composition and prey intake rates of Australian magpies in a New Zealand population were investigated to determine whether flocking, non-territorial birds were less successful predators of pasture invertebrates than territorial birds. Time-activity budgets showed that flock magpies foraged throughout the day during February and March, indicative of a prey shortage at that time. Prey abundance was therefore measured in the autumn months but there was no evidence of a large quantitative difference in the food supply of flock and territorial magpies. Significantly fewer invertebrates were collected from pitfall traps in the flock foraging area, but these were counterbalanced by significantly more scarab beetle larvae and more flying insects, obtained by sweep-netting. When compared with territorial magpies, flock birds exhibited similar peck rates, prey intake rates, and earthworm intake rates. While there were qualitative differences in the diets of flock and territorial magpies, determined by faecal analysis, five of seven prey items analysed were captured in the same relative proportions by the flock and territorial birds. Earthworms exceeded their relative availability in the diets of flock birds, and more scarab larvae were found in the diets of territorial magpies. Non-territorial magpies were therefore no less successful than territorial birds during the autumn, and it is suggested that the function of flocking behaviour may eventually be discovered through consideration of why non-territorial birds never occur as solitary individuals, rather than emphasizing comparisons with territorial magpies.     

Water loss from Jackass Penguin Spheniscus demersus eggs during natural incubation

The water budget of incubating Jackass Penguin eggs was studied on Marcus Island, South Africa, and complementary measurements were made in the laboratory. The mean ambient temperature was 16-5 "C and the mean humidity was 12-4 Torr (89% relative humidity). The temperature of incubated live and water-filled eggs ranged between 14^oCand 37 ^oC. The mean calculated egg temperature was 34-9' C. The mean brood patch temperature was 37-1 ^oC, slightly lower than the cloacal temperature (37.8 ^oC). The mean brood patch area was about 38 cm². The rate of water loss was 411 mg day^-1. The total diffusive water loss during 37 days of incubation was, as predicted, 15-2% of the initial 100-3 g egg mass. The total pore number was 6245 per egg and the shell thickness was 577 fira. It is suggested that the eggshell parameters, incubation length and nesting behaviour are compensated in such a way that an egg-to-nest water vapour pressure difference lower than commonly found is sufficient to bring about the normal total water loss.     

Winter energy requirements of soricine shrews

The energy requirements of shrews under snow cover have not been determined. This is an expository paper attempting to ascertain the daily energy budget of a soricine in subnivean conditions. By means of extrapolation from past work (Morrison, Ryser & Dawe, 1959; Gebczynski, 1965, 1971), a winter individual of Sorex cinereus (mean weight of 2–6 g) in southern Manitoba has a rather high energy budget at 2^oC of about 38-6 kj or 9-2kcal per day. It is assumed that winter prey include invertebrates and small mammals, and their biomass and energy content are estimated. Compensatory factors reducing energy needs are considered, i.e. Dehnel's Phenomenon, extended periods in the nest, low population densities and adequate snow cover. We can probably reduce the above figure for a daily energy budget when considering all the above factors, so that about 30 kj are required daily at the mean subnivean temperature of – 4–5^oC.     

BIOLOGY OF THE BANK CORMORANT,PART 3: FORAGING BEHAVIOUR

Cooper, J. 1985. Biology of the Bank Cormorant, Part 3: Foraging behaviour. Ostrich 56: 86–95.

The Bank Cormorant Phalacrocorax neglectus, endemic to southern Africa, is primarily a solitary inshore forager. Bank Cormorants forage Primarily on the bottom among kelp beds but also may forage over shingle or coarse sand substrates or in midwater. Breeding birds forage up to 9 km from their colony. Little is known of foraging depth but birds may dive as deep as 28 m. Mean dive duration was 44,9 s and ratio of dives to surface rests was 2,18. In most cases prey is swallowed under water, presumably to avoid kleptoparasitism. Bank Cormorants foraged during daylight hours from before sunrise to after sunset. Birds did not forage in exceptionally rough seas. Mean female foraging bout duration (84,3 min) was significantly longer than that of males (68,4 min) in breeding individuals. Breeding males undertook significantly more foraging bouts (3,47 boutdday) than did females (3,02 bouts/day). No significant differences were found between the sexes when total time spent foraging/day by breeding birds was compared. It is not clear why males foraged more often, but for shorter periods, than did females, but the differences may be related to sexual dimorphism, males being larger than females.     

The novel ‘tongue‐bite apparatus’ in the knifefish family Notopteridae (Teleostei: Osteoglossomorpha): are kinematic patterns conserved within a clade?

Osteoglossomorph fishes are unique in possessing a specialized feeding mechanism, the tongue-bite apparatus (TBA) involving the hyoid apparatus. The TBA is associated with two unique behaviour patterns - raking and open-mouth chewing - used to disable and macerate prey. The kinematics of these two behaviours was compared in two species of knifefishes (family Notopteridae): Xenomystus nigri (Gunther, 1868) and Chitala ornata (Gray, 1831) using high-speed video (250 frames s"¹). Both univariate and multivariate analyses indicated that there were significant interspecific differences in both raking and open-mouth chewing. Raking can be divided into three stages; the preparatory phase, power stroke, and recovery phase. During the power stroke posterior motion of the pectoral girdle and neurocranial elevation both appear to play a major role in prey reduction. In Xenomystus the power stroke involves significantly greater levels of neurocranial elevation (35^o) and pectoral girdle motion (38% of head length; 9.5^o) than that found in Chitala (neurocranial elevation 11^o; pectoral girdle motion 11% of head length and 5^o). Indeed, Xenomystus represents the most extreme raking behaviour of any osteoglossomorph thus far studied. Temporal displacement variables demonstrated that the power stroke in Xenomystus is significantly faster than in Chitala. Although some of the interspecific differences might be size related, these data suggest that a greater degree of difference exists in these highly specialized behaviours than previous work has demonstrated. These findings support the notion that biomechanical duplication (an additional ligament found in osteoglossomorphs) could be linked to increased functional versatility.     

Territorial behavior of Western Sandpipers on their nonbreeding grounds: effect of sex and foraging interference

ABSTRACT Nonbreeding shorebirds often alternate social structure between anonymous flocks and territorial behavior in response to different environmental factors. To evaluate specific drivers for one species, we studied the spacing behavior of wintering Western Sandpipers (Calidris mauri) at Bahía Santa María, northwestern Mexico, using behavioral observations. The density, population structure, and territorial behavior of Western Sandpipers differed among three habitat types. Cattail marshes supported 110 birds per ha, 49% males, and no territorial birds. Mangrove flats supported 288 birds per ha, 58% males, and 5% territorial birds. Brackish flats supported 365 birds per ha, 76% males, and 7% territorial birds. Territories consisted of rectangular strips (5–9 m long, N= 77). Territory length was not related to either bird density or number of territorial birds by plot, but was positively related to nearest bird distance. Aggression rate was inversely related to territory length, suggesting that territory length is set by the costs of defense. Foraging rate was independent of territory length, and prey densities in territories did not differ from those in areas used by nonterritorial birds. Males were more likely to be territorial and had a higher aggression rate than females, suggesting that males, which forage more on surface prey, were more affected by foraging interference. Our results suggest that the territorial behavior of Western Sandpipers in our study was an opportunistic strategy to reduce foraging interference. The variation in spacing behavior we documented provides evidence that interference competition affects the social structure of Western Sandpipers during the nonbreeding season.     

The effect of sward height and drainage on Common Starlings Sturnus vulgaris and Northern Lapwings Vanellus vanellus foraging in grassland habitats

Agricultural change is often cited as a causal factor in the decline of the UK's farmland birds because bird declines have mirrored changes in agricultural practices. Although much is known about the mechanisms driving population declines on arable systems, mechanisms in grassland systems are relatively poorly studied, despite receiving a similar degree of intensification. Agricultural intensification may affect bird declines by reducing food abundance or accessibility, forager mobility or predation risk. Here we examine experimentally the effects of sward height on the foraging behaviour of adult Common Starlings Sturnus vulgaris , and the effects of sward height and drainage on the behaviour of Northern Lapwing Vanellus vanellus chicks. Both species are declining across the UK and both forage in farmed grassland habitats, but they differ in their foraging methods because Starlings probe for soil invertebrates whereas Lapwing chicks glean prey from surfaces. Overall, after controlling for prey abundance, short swards were found to be more productive for both species. Prey capture rate within foraging bouts did not differ with sward height for Starlings, but Starlings spent more time foraging on short swards and captured 33.2% more prey. Starlings walked more steps on short swards. Lapwing chick foraging rates declined as sward height increased. Soil moisture was not found to be a predictor of Lapwing chick foraging rates within the observed range. Our results suggest that short swards are a more profitable foraging habitat for soil and surface invertebrate feeders. Short swards may facilitate surface prey detection, improve forager mobility and increase foraging time by altering vigilance patterns. Provision of short swards in areas where these are lacking could be simple method of improving foraging habitats for grassland birds.     

Seasonal Variations in the Diet and Foraging Behaviour of Dunlins Calidris alpina in a South European Estuary: Improved Feeding Conditions for Northward Migrants

During the annual cycle, migratory waders may face strikingly different feeding conditions as they move between breeding areas and wintering grounds. Thus, it is of crucial importance that they rapidly adjust their behaviour and diet to benefit from peaks of prey abundance, in particular during migration, when they need to accumulate energy at a fast pace. In this study, we compared foraging behaviour and diet of wintering and northward migrating dunlins in the Tagus estuary, Portugal, by video-recording foraging birds and analysing their droppings. We also estimated energy intake rates and analysed variations in prey availability, including those that were active at the sediment surface. Wintering and northward migrating dunlins showed clearly different foraging behaviour and diet. In winter, birds predominantly adopted a tactile foraging technique (probing), mainly used to search for small buried bivalves, with some visual surface pecking to collect gastropods and crop bivalve siphons. Contrastingly, in spring dunlins generally used a visual foraging strategy, mostly to consume worms, but also bivalve siphons and shrimps. From winter to spring, we found a marked increase both in the biomass of invertebrate prey in the sediment and in the surface activity of worms and siphons. The combination of these two factors, together with the availability of shrimps in spring, most likely explains the changes in the diet and foraging behaviour of dunlins. Northward migrating birds took advantage from the improved feeding conditions in spring, achieving 65% higher energy intake rates as compared with wintering birds. Building on these results and on known daily activity budgets for this species, our results suggest that Tagus estuary provides high-quality feeding conditions for birds during their stopovers, enabling high fattening rates. These findings show that this large wetland plays a key role as a stopover site for migratory waders within the East Atlantic Flyway.     

The impact of predation by birds on bat populations in the British Isles

The role played by predation of birds in the mortality of British bats is assessed. A review of dietary studies and anecdotal accounts revealed eleven species of bird which occasionally feed on bats–Herring Gull Larus argentatus , Black-headed Gull Larus ridibundus , Rook Corvus frugilegus , Carrion Crow Corvus corone , Little Owl Athene noctua , Short-eared Owl Asio flammeus , Kestrel Falco tinnunculus , Hobby Falco subbuteo , Merlin Falco columbarius , Peregrine Falco peregrinus and Sparrowhawk Accipiter nisus. A further three species feed more frequently on bats–Barn Owl Tyto alba , Tawny Owl Strix aluco and Long-eared Owl Asio otus. Rates of predation were very low accounting for only 00034^o_o of items taken by small hawks and falcons ( n items = 29 519) but 0035^o, of prey taken by owls ( n items = 99 479). By multiplying together the average annual prey capture rates of the predators, assessed from their energetic food requirements and direct observations of prey intake rates, the British populations of the predators and the contribution to the diet made by bats, the annual number of bats which die each year as a result of predation was estimated. The total losses of bats to predation might amount to about 201 400 bats/annum. The most significant predators are Tawny Owl ( c. 168 850 bats/annum), Barn Owl (c. 8800 bats/ annum), Long-eared Owl (c. 10 200 bats/annum) and Kestrels (c. 8400 bats/annum). This predation by avian predators would account for about 11 % of the annual mortality of British bats. An assessment of the biases involved in this calculation suggests it is probably a minimum estimate. Despite the apparent low representation of bats in the diets of predatory birds, the effects of this predation on bat behaviour and population dynamics cannot be ignored.     

Day and night feeding territoriality in Willets Catoptrophorus semipalmatus and Whimbrel Numenius phaeopus during the non-breeding season in the tropics

This study was carried out to determine whether or not, and under what conditions, Willets Catoptrophorus semipalmatus and Whimbrel Numenius phaeopus continue to defend and use their daytime feeding territories at night in a tropical environment. The study was conducted in coastal Venezuela by registering, by night and by day, the behaviour and the position of colour-marked and radio-tagged birds. Night observations were made with the use of a light intensifier.
In Willets, both territorial and non-territorial birds were observed. The use and defence of territories was observed only on sandy mud areas where Fiddler Crabs Uca cumulanta were plentiful; the soft mud sites were used only by non-territorial birds. All Whimbrel were territorial. Individuals of both species defending a given space during daytime continued to occupy and defend the same area during the night. The proportion of birds foraging at night was higher on moonlit than on moonless nights. Territorial defence involved alert postures, parallel walks, calls and the chasing of intruders. Agonistic encounters between Willets and Whimbrel and other species were rare. Willets were territorial at all states of the tide except when high tides flooded the territories. Tide had no effect on the time of feeding in Whimbrel. Moonlight was clearly the factor conditioning the occurrence of nocturnal foraging on territories by both species. Foraging strategies and the type of prey and substrata explain why the incidence of night foraging varied with moonlight in territorial Willets and Whimbrels and not in non-territorial individuals.     

Calling behaviour in M.convecta females under different temperature and photoperiodic conditions

Abstract. . The effects of temperature and photoperiod on calling behaviour in females of the Australian common armyworm, Mythimna corwecta (Walker) (Lepidoptera: Noctuidae), were studied in the laboratory. Age at first calling was greater at 15 and 20^oC compared with 25^oC, but there were no significant differences between 25 and 30^oC. Females kept at 10^oC did not call, but if transferred at 10 days to 20^oC they called after a period similar to the pre-calling period at constant 20^oC. Onset time of calling was earlier at 15 and 20^oC compared with 25 and 30^oC, but there were no significant differences in calling duration. At 20^oC, age at first calling was increased by shorter day length (12h and 14h, compared with 16h), and there was an interaction between daylength and moth age affecting the duration of calling. Onset times advanced with longer daylength, but peak calling was later in relation to the lengtii of the scotophase. These results are discussed in connection with migration in M.convecta. Evidence for a circadian rhythm of calling was found in females entrained for 3 and 8 days after emergence under reverse-cycle conditions and then transferred to constant darkness. However, after 56 h and 80 h respectively of darkness, calling became de-synchronized. Subsequently, it appeared to re-synchronize to a different clock, which approximately correlated with the normal photoperiods the moths had experienced during larval development.     

Germination ecology of drupelets of the fig (Ficus carica L.)

Abiotic and biotic factors and their effects on germination of fig drupelets were studied. The drupelets germinated between 10^oC and 30^oC.Constant humidity was necessary for germination and frequent drying out of the substrate was unfavourable. Total darkness for the whole duration of the experiments had a slighly negative effect on germination. The fastest germination occurred at constant humidity and an alternating temperature of 20/30^oC with exposure to light for 8 hours (at the higher temperature) and to darkness for 16 hours (at the lower temperature).Natural or artificial removal of drupelets from the syconium guaranteed a high germination percentage, whereas no germination occurred in drupelets left inside the syconium. Hence birds and mammals act as dispersal agents and mediators of germination. As they eat pieces of fig syconium, they free the drupelets from the flesh, eliminating the effect of inhibitors and/or microenvironments with high osmotic pressure inside the syconium. These findings support the hypothesis that germination occurs in autumn or spring depending on the climatic zone in which the species grows.     

Sibling cannibalism among juvenile vundu under controlled conditions. I. Cannibalistic behaviour, prey selection and prey size selectivity

Sibling cannibalism among vundu Heterobranchus longifilis larvae started at the age of 4 days, with the prey caught tail-first then swallowed up to the head, which was eventually discarded (type I cannibalism). At 17 days old, this type of cannibalism vanished and was replaced by the ingestion of the whole prey (type II cannibalism), which could only be exerted by predators six times as heavy as their prey. Type II cannibalism consisted of a seemingly opportunistic ambush attack by a formerly passive predator towards a disorientated prey. It required no preliminary aggression or chase, or even contact with the prey, suggesting that the attack was not mediated by the tactile sense, and that cannibalism was independent of aggressive behaviour. When alternative food resources (formulated feed, live tilapia prey) were available, the intensity of cannibalism decreased but pellet-eaters or tilapia predators always achieved lower growth rates than those feeding on conspecifics, suggesting that cannibalism was the most advantageous foraging tactic. Losses to cannibalism among populations of 30-day old juvenile vundu with an initial ratio of 4% of cannibals were as high as 75·5–79·9% over 15 days. Predation peaked during the first days (up to 2·8 prey C⁻¹ day⁻¹), then vanished progressively as surviving prey grew faster than cannibals and escaped their predation. Cannibals preferred consuming the largest prey available with respect to the logistics of cannibalism (body weight ratio of 6·0). This preference for large prey was interpreted both as a foraging tactic aiming to maximize the energetic return, and as foraging strategy enabling the cannibals to exploit their prey as long as possible. Based on these data, comprehensive models of the impact of cannibalism on vundu populations were developed.     

Thermal response and reversibility of diapause in the eggs of Locusta migratoria

Abstract. Eggs of a local population of Locusta migratoria L. (Orthoptera: Acrididae) collected near Hirosaki (40.5^oN) entered diapause when incubated at temperatures between 20 and 35^oC. For diapause development the optimum temperature was 10^oC. The lower thermal threshold for post-diapause development was 14.7^oC. After chilling at 10^oC for 20 days, the rate of hatching varied with incubation temperature, being 0, 61% and 81% at 20, 25 and 30^oC, respectively. After chilling for 40 days or more, however, almost all eggs hatched at 20–30^oC. Diapause with a reduced intensity seemed to be eliminated easily by a high temperature of 25 or 30^oC.
When eggs chilled at 10^oC for 20 days were kept at 20^oC for 7 days or more before incubation at 25^oC, almost all eggs maintained diapause. Most eggs chilled at 10^oC for three 10 day periods separated by 3 days of warming at 25^oC failed to terminate diapause. Daily alternations of 10^oC (18 h) and 25^oC (6h) decreased the diapause-terminating effect of chilling. These facts suggest that diapause intensity can be restored if eggs chilled insufficiently are exposed to a moderately high temperature. This reversible change in diapause intensity would play an important role in maintaining diapause before winter.     

The Relative Importance of Habitat Structure and of Prey Characteristics for the Foraging Success of Water Pipits (Anthus spinoletta)*

While many studies on foraging have related energy gain to the density and the size of prey, only few have investigated whether and how habitat structure modifies the gain through affecting foraging success. In this study, the influences of habitat structure and prey characteristics on the foraging success of water pipits, Anthus spinoletta, were investigated experimentally. The birds take longer to find prey in tall than in short vegetation. The effects of vegetation on searching times differ between prey types. These differences are probably caused by variation in prey behaviour and in cryptic colouration, but not by prey size. Searching times increase with decreasing density for mealworms and tipulids, but not for caterpillars. Handling large prey items requires more time than handling smaller prey. Tipulids and caterpillars, which were offered alive, are handled for a longer time than dead mealworms of corresponding size. The success of attacks on flying insects is probably influenced by the prey's flight speed: fast houseflies are missed more often than slow tipulids. Overall, the results show that the time costs of foraging water pipits are influenced to a comparable degree by vegetation structure, by prey density and by other specific prey characteristics such as camouflage, hiding behaviour or agility. The amount of food gathered per unit time is determined primarily by factors that affect searching times, and less by handling and travelling times. Insertion of our data into an optimal diet model leads to the prediction that water pipits should be generalist foragers, which agrees with the observed behaviour.     

Foraging behaviour of Lesser Sheathbills Chionis minor exploiting invertebrates on a sub-Antarctic island

Summary During winter (May through October) many Lesser Sheathbills Chionis minor at Marion Island in the sub-Antarctic were obliged to leave their preferred foraging habitat in penguin colonies to forage for invertebrates on the island's coastal plain. The study describes factors affecting feeding success, time budgets and predation risk of the sheathbills which exploited these small, patchily dispersed prey. The birds appeared to select prey 1 mm in diameter, and ignore smaller, common invertebrates.Sheathbills were highly selective of foraging habitat. During 17 censuses made through the winter, 97% of the 1,504 birdsightings were at only eight of the 19 available vegetation types. Multiple regression analysis revealed that prey density was the most important criterion in habitat preference, followed by plant canopy height and distance of the habitat from the sea. These variables accounted for 78% of the variance of habitat use. Focal-animal observations in a sample of habitats showed that feeding success was correlated with prey density and distance from the sea. Tall vegetation impeded the locomotion and foraging of sheathbills. The sheathbills reduced predation risk from skuas Catharacta lonnbergi and travelling time by foraging near the shore. The spatial distribution of prey within vegetation types was apparently unimportant in habitat selection.During winter 83% of the sheathbills in the study foraged communally and 98% roosted communally. Flocks occurred only on good quality habitat and flocking probably facilitated habitat selection. Feeding success increased initially with increasing flock size but decreased in flocks greater than 15 birds, which was attributed to localized prey deletions. The sheathbills spent 88% of the daytime foraging; and feeding, looking around and walking comprised 99% of foraging time. Feeding time increased with increasing flock size, looking around decreased but walking was unaffected. Aggression was rare, was unaffected by flock size and did not significantly affect feeding. A probability model showed that sheathbills could greatly reduce predation risk by flocking but the benefits would not improve much in flocks greater than eight birds.The habitat selection, time budgets and feeding success of adults, subadults and juveniles were very similar.The exploitation of terrestrial invertebrates by sheathbills was interpreted as an expansion of the population's trophic niche to tap an underexploited resource on a species-poor island.     

Cold shock and heat shock: a comparison of the protection generated by brief pretreatment at less severe temperatures

Abstract Brief exposure to low (0^oC) or high (40^oC) temperature elicits a protective response that prevents injury when the flesh fly, Sarcophaga crassipalpis Macquart, is subjected to more severe cold (-10^oC) or heat (45^oC). Both the low and high temperature responses were found in all developmental stages of the fly, but were most pronounced in the pupal and pharate adult stages. The protective responses generated by brief exposure to 0 or 40^oC appear similar in that both result in a rapid acquisition of cold or heat tolerance and a loss of protection after the flies are returned to 25^oC. The protection generated by chilling is obvious within 10 min of exposure to 0^oC while a 30 min exposure to 40^oC is required to induce the high temperature protection. High temperature protects against cold shock injury within a narrow range (around 36^oC) but we have no evidence that low temperature can protect against heat injury. We previously demonstrated that the rapid increase in cold tolerance correlates with concomitant increases in glycerol concentration, but in this study we found no significant elevation in glycerol in heat-shocked flies. Thus the physiological and biochemical bases for the rapid responses to cold and heat appear to be different.     

Effects of small-bodied fish on invertebrate prey and foraging patterns of waterbirds in Aspen Parkland wetlands

Competition between large-bodied fish and waterbirds for aquatic invertebrates is well documented in oligotrophic lakes. Recent evidence suggests that small-bodied fish that colonize eutrophic, hypoxia-prone wetlands such as prairie potholes can also reduce aquatic invertebrates, but the effects of these reductions on breeding waterbirds have so far not been directly documented. We added brook stickleback (Culaea inconstans) and fathead minnow (Pimephales promelas) to a fishless wetland in Aspen Parkland potholes in central Alberta, Canada. We monitored invertebrate biomasses and the foraging effort of blue-winged teal (Anas discors) and red-necked grebe (Podiceps grisegena) before and after the addition, relative to reference wetlands with and without fish. Fish reduced the biomass of gastropod prey of blue-winged teal, and teals increased foraging effort when fish were added. When the fish failed to overwinter due to hypoxic conditions, gastropod biomass increased, but teal foraging effort did not return to pre-treatment levels. Amphipods and chironomids increased following fish addition, possibly due to indirect positive effects of fish. Red-necked grebes did not exhibit any changes in foraging effort as a result of the fish addition or the subsequent fish extirpation. Grebes in Aspen Parkland appear to treat fish and invertebrates as equivalent prey. This study suggests that small-bodied fish in eutrophic systems can reduce some important invertebrate prey and change foraging behaviour of blue-winged teal and other waterbirds that rely on those invertebrates. Land-use practices that encourage survival of colonizing fish through drought years in Aspen Parkland wetlands, such as wetland consolidation, should not be encouraged.     

DIET AND DISPERSION OF THE GREAT ARGUS ARGUSIANUS ARGUS

The food and dispersion of Great Argus were studied in two West Malaysian forested areas from 1975 to 1978. Invertebrates on the litter surface lived at low density and were typically large and solitary; those beneath the litter were abundant but typically tiny and colonial. Great Argus feed mainly on fallen fruit but rely for protein on the large, solitary litter-surface invertebrates and were never seen scratching for food. Adult males maintain small, discrete home ranges in which they forage alone. Much of their time is spent resting, which reduces energy expenditure. Females also forage alone and may be similarly inactive. Great Argus are very large and their requisite large invertebrate prey cannot be obtained beneath the litter; scratching has been eliminated as a foraging technique. A strategy of litter-surface searching necessitates solitariness, to avoid intraspecific feeding competition. The large size of Great Argus is related to a low rate of energy turnover and energy conservation by prolonged resting. Both sexes are probably territorial during the breeding season and larger territories then may limit population density.     

Hourglass and oscillator expressions of photoperiodic diapause response in the cabbage moth Mamestra brassicae

Abstract. Both oscillator and hourglass features are found in the photoperiodic response that controls the pupal winter diapause of Mamestra brassicae. The expression of oscillatory response to extended long-night cycles is temperature dependent, i.e. circadian resonance appears at 23 and 25^oC but not at 20 and 28^oC. At 20^oC, scanning of extended scotophases by a short light pulse does not reveal any clear circadian rhythmicity. However, a circadian feature of the photoperiodic response is indicated even at 20^oC by a bistability phenomenon, i.e. either one of the two dark periods in symmetrical skeleton photoperiods determines the diapause response depending on the phase angle with the preceding (entraining) light-dark cycles. At 20 and 25^oC, the incidence of diapause increases as a function of the number of light–dark cycles regardless of the cycle length (T) , if T is 24 h or 2 X 24h (with a 12 h light period). A non-diel cycle (r=36h) is less effective, suggesting that disturbance of the circadian organization partly impairs the diapause-inducing function. The inductive effect of a long night is largely affected by temperature, and becomes saturated with eight cycles at 20^oC and 14 cycles at 25^oC. Presumably, an hourglass mechanism measures the dark time, and a circadian component involved in some later sequence of the photoperiodic response may or may not be expressed depending on the mode of interaction between them.