Adaptive variation in the foraging behaviour of Grey Plover Pluvialis squatarola and Whimbrel Numenius phaeopus

Between-site variation in the diet and foraging behaviour of a fixed-method forager, the Grey Plover Pluvialis squatarola , and a versatile forager, the Whimbrel Numenius phaeopus , was examined and compared at nine tropical and south temperate sites. Grey Plover always foraged in a run-stop-search manner but changed the emphasis of components of this behaviour in response to prey type. Whimbrel foraged tactilely or visually and readily changed foraging speed and habitat. Both species ate a variety of prey species depending on their availability. The diet of Grey Plover was dominated by small prey, polychaetes and crabs, and Whimbrel ate mostly crabs, with Whimbrel being the more specialized in prey choice. The broader diversity of prey types consumed by Grey Plover was attributed to limitations imposed by obligate visual foraging, whereas the flexible foraging behaviour of Whimbrel allowed the latter to concentrate their efforts on the most profitable prey. Grey Plover appeared to have a density upper limit, determined by their stereotyped foraging behaviour. Whimbrel densities varied greatly in response to prey type and foraging method. Foraging effort could not be predicted from measurements of instantaneous daytime energy intake rates, and reasons for the lack of this relationship are discussed.     

Duration of primary moult affects primary quality in Grey Plovers Pluvialis squatarola

Feather wear is the natural degradation and breakage of feather structure during the interval between moults. Different rates of feather wear have been observed for primaries of free-living populations of several species of passerines and waders, and this variability has been linked to different concentrations of melanins. In this study primary moult duration explained 59% of the variation in annual rates of primary abrasion (percentage wing length loss) of seven Grey Plover wintering populations, while migration distance explained 14%. The analysis suggests that primary moult duration plays a key role in determining primary durability and hence primary quality. Long distance migrants might evolve more durable primaries, despite the higher predation risks and energetic costs of a prolonged moult. Partial or complete pre-breeding primary moults of first-year waders and complete biannual moults of some passerines might have evolved under selective forces favouring migration with unabraded primaries.     

Biometrics,possible breeding origins and migration routes of South African Grey Plovers,Pluvialis squatarola

Data on 355 Grey Plovers, Pluvialis squatarola, ringed in South Africa in 1971–1997 were analysed. Biometrics could not identify well-defined origins in the Siberian breeding grounds, but suggested the presence of birds from east of the Gydan Peninsula. Comparisons with populations spending the non-breeding period elsewhere did not allow a clear definition of the flyways used, although ringing recoveries indicated a migration route crossing the Mediterranean/Black Sea region during both southward and northward movements. First-year birds were on average smaller than adults. Bill length increased during the first year, reaching adult length by June. First-year wing length decreased by 4% before the outermost primary was shed, adult wing length did not change with season.     

Primary moult, body mass and migration of Grey Plovers Pluvialis squatarola in Britain

Long-distance migrants have evolved complex strategies for the timing of their annual moult, fattening and migration cycles. These strategies are likely to vary at different stages of a bird's life. Ringing data on 6079 Grey Plovers Pluvialis squatarola , caught on the Wash, England, between 1959 and 1996, were analysed to relate migratory strategies to patterns of primary moult and body mass changes. Adults returning from breeding grounds had a shorter and delayed primary moult (duration 90 days, starting date 19 August) in comparison with over-summering birds (duration 109 days, starting date 5 June). Three categories of migrant adults were identified on the basis of primary moult and body mass: (1) birds which did not moult, but increased body mass and migrated further south; (2) birds which moulted 1–3 inner primaries, suspended moult, increased body mass and migrated; and (3) birds which completed or suspended moult and wintered locally. In birds of the second category, timing of primary moult and body mass increase overlapped. Among wintering birds, 38% were in suspended moult. Ninety-six per cent of birds that suspended moult at the beginning of winter were males and almost all completed moult in spring. Grey Plovers which left Britain in autumn had an average body mass of 280 g, enough to reach southern Morocco without refuelling. Both wintering adults and first-year birds showed a prewinter body mass increase, peaking in December. Adults had a synchronized premigratory body mass increase in May, which suggested a negligible presence of African migrants. The average departure mass for spring migration, estimated at 316 g, would allow birds to fly non-stop to the Siberian breeding grounds in western Taymyr.     

Primary moult,biometrics, mass and age composition of Grey Plovers Pluvialis squatarola in southeastern India

Little is known about the biology of waders wintering in southern Asia; this paper deals with the Grey Plover Pluvialis squatarola, a species extensively studied only in western Europe. Adult Grey Plovers wintering in southeastern India underwent primary moult in autumn; the duration was estimated to be 127 days, with mean starting date 1 September and mean completion date 5 January. Some first-year Grey Plovers initiated primary moult in late winter and spring, and completed this moult the following spring. The average mass of adults on arrival in September was 200 g, fluctuated close to 220 g from October to February, and increased to 280 g near the end of May. The mass variation did not show the January peak observed in western Europe. Breeding productivity, measured as the percentage of first-year birds in winter catches, varied between 5% and 70% over six years, and showed a positive correlation with that of Dark-bellied Brent Geese Branta b. bernicla in western Europe and Curlew Sandpipers Calidris ferruginea in South Africa.     

Contrasts in energy intake and expenditure in sit-and-wait and widely foraging lizards

Summary Daily energy metabolism and water flux were measured with doubly labeled water in the free-living insectivorous lizards Cnemidophorus tigris (mean body mass 15.7 g) and Callisaurus draconoides (8.6 g) in June 1979 in the Colorado Desert of California. C. tigris was an active forager; it spent 91% of its 5-h daily activity period in movement. C. draconoides was a sit-and-wait predator; it spent less than 2% of its 10-h activity period in movement. C. tigris had significantly higher rates of field energy metabolism and water influx (210 Jg^-1 day^-1, 36.8 ul H₂O g^-1 day^-1, N=19) than C. draconoides (136, 17.1, N=18). There were no significant differences between the sexes within either species.The extra costs of free existence were calculated from differences between field metabolic rates and maintenance costs estimated from laboratory respirometry. Rates of energy metabolism during the field activity period were about 1.5x resting levels at 40° C (field active body temperature) for C. draconoides and 3.3 x resting levels at 40° C (field active body temperature) for the more active C. tigris. Feeding rates calculated from water influx data were 13.3 mg g^-1 day^-1 for C. tigris and 5.8 mg g^-1 day^-1 for C. draconoides. Though C. tigris had a high rate of energy expenditure, its foraging efficiency was higher than C. draconoides'.     

Influence of sward structure on daily intake and foraging behaviour by horses

The spatial heterogeneity of grasslands determines the abundance and quality of food resources for grazing animals. As plants mature, they increase in mass, which allows greater instantaneous intake rates, but the cell wall concentrations increase too, reducing diet quality. In ruminants, daily intake rates are often constrained by the time needed for the ingesta to pass through the rumen, which is influenced by the rate of digestion. It has been suggested that the digestive constraint should have much less effect on hindgut fermenters such as equids. Horses play an increasing role in the management of grasslands in Europe, but the data on the influence of the heterogeneity of the vegetation on their daily intake and foraging behaviour are sparse. We report here the results of a preliminary study concerning the effects of sward structure on nutrient assimilation and the use of patches of different heights by horses grazing successively a short immature, a tall mature and a heterogeneous pastures (with short and tall swards). Daily nutrient assimilation was higher in the heterogeneous pasture compared to the short (+35%) and the tall (+55%) ones. The digestive constraints may have limited voluntary intake by horses on the tall swards. In the heterogeneous pasture, the mean height used for feeding (6 to 7 cm) by horses was intermediate between the heights used in the short (4 to 5 cm) and tall pastures (22 to 23 cm), and the animals may thus have benefited from both short swards of high quality and tall swards offering a higher instantaneous intake rate.     

Seasonal and diurnal variation in vocal behaviour of the South Island robin

Abstract

The proportion of time spent by South Island robins (Petroica australis australis) of different sex, age, and status giving full song, subsong, and downscales, on a monthly and diurnal basis, is described. Females never gave full song. Males did so throughout the year, but least while moulting. Since bachelors sang significantly more than paired males, song probably functioned largely to attract females. Breeding males sang most during the pre-laying stage. From January to April adult males sang significantly more than immature males, but from May to July the reverse was true. Robins sang mainly in the early morning. Robins generally spent less than 0.5% of time during any stage of the breeding cycle giving sub-song. Outside the breeding season, males spent significantly more time giving sub-song than females, and adult males sang more than immature males. Robins in all categories gave more sub-song during the moult than afterwards. The diurnal pattern of sub-song production was bimodal. About half the downscales were followed by no obvious action from the caller or response from other robins, nor were the calls preceded by an obvious event. About 25% of the downscales were associated with interactions between members of a pair; the remainder were associated with territorial defence. Downscales probably served to maintain contact between members of a pair and to advertise possession of territory. These calls were heard rarely in the breeding season, and most frequently in April. Males gave downscales more than females, but use of downscales by both sexes was evenly distributed through the day.     

The relationship between foraging behaviour and energy expenditure in Antarctic fur seals

By using time-depth recorders to measure diving activity and the doubly-labelled water method to determine energy expenditure, the relationship between foraging behaviour and energy expenditure was investigated in nine Antarctic fur seal females rearing pups. At-sea metabolic rate (MR) (mean of 6.34 ± 0.4 W. kg^-1; 4.6 times predicted BMR) was positively correlated to foraging trip duration (mean of 4.21 ± 0.54 days; r²= 0.5, P < 0.04). There were no relationships between MR and the total number of dives, the total time spent diving or the total vertical distance travelled during the foraging trip. There was, however, a close negative sigmoidal relationship (r²= 0.93) between at-sea MR and the proportion of time at sea spent diving. This measure of diving behaviour may provide a useful, inexpensive means of estimating foraging energy expenditure in this species and possibly in other otariids. The rate of diving (m.h^-1) was also negatively related to at-sea MR (r²= 0.69, P < 0.005). Body mass gain during a foraging trip had a positive relationship to the time spent at sea (r²= 0.58, P < 0.02) and the total amount of energy expended while at sea (r²= 0.72, P < 0.004) such that, while females undertaking long trips have higher metabolic rates, the energetic efficiency with which females gain mass is independent of the time spent at sea. Therefore, within the range of conditions observed, there is no apparent energetic advantage for females in undertaking foraging trips of any particular duration.     

Opposite effects of tryptophan intake on motor activity in ring doves (diurnal) and rats (nocturnal)

The role of l-tryptophan as precursor of serotonin and melatonin synthesis on activity-rest rhythm was studied in ring doves, Streptopelia risoria, as a representative of diurnal animals and rats, Rattus norvegicus, as a typical nocturnal one. The animals were housed in cages equipped for horizontal activity recording in a thermostatized chamber and submitted to a 12/12h light/dark photoperiod (lights on at 08:00 h). After acclimatization, the animals received vehicle (methylcellulose) and l-tryptophan (240 mg/kg) by esophagic cannula 2h before the onset of either light or dark phase. Also, oral melatonin (2.5mg/kg) was tested for comparative purposes. After nocturnal l-tryptophan administration, rats showed increased activity (149%), while the opposite occurred in ring doves (39% decrease). No significant changes were found after diurnal l-tryptophan intake in either species. Melatonin produced effects similar to those of l-tryptophan. These results suggest that the effects of l-tryptophan administration are dependent on the nocturnal/diurnal habits of the studied species and, most probably, are mediated by increased melatonin synthesis.     

Faunal assemblage composition and paleoenvironment of Plovers Lake, a Middle Stone Age locality in Gauteng Province, South Africa

Plovers Lake is a dolomitic cave infill located approximately 45 km northwest of Johannesburg in the Bloubank Valley, Gauteng Province, South Africa. Excavations between 2002–2004 revealed a rich and diverse fauna, a moderate-sized stone tool assemblage of Middle Stone Age (MSA) character, and human skeletal remains. Two principal depositional units are recognized: 1) a disturbed ex situ component that was likely displaced from 2) an otherwise relatively undisturbed in situ component from which the human skeletal material was recovered. The in situ depositional unit is bracketed by 2 flowstone layers, with U-series dates of 62.9 (± 1.3) ka for the capping flowstone and 88.7 (±1.6) ka for the underlying flowstone. A single isochron ESR date of 75.6 (± 5.6) ka corroborates the U-series dates. This paper presents an analysis of the mammalian, bird, and reptile faunas recovered from these two units. The two faunal assemblages show close correspondence in taphonomic, taxonomic, and ecological composition, supporting a common origin for both the ex situ and in situ components. Although human skeletal material, cut-marked bone, and stone tools have been recovered, these indications are too rare to consider Plovers Lake a human occupation site. Instead, a high abundance of carnivores, coprolites, and carnivore damaged bones point to brown hyenas as the principal, though not exclusive, bone accumulating agent. In the absence of a significant taphonomic bias relating to accumulating agent, Plovers Lake allows us to document an environment occupied by MSA humans, even if the humans were not resident in the cave itself. We reconstruct the paleoenvironment of Plovers Lake as predominantly grassland, though it was colder, moister, and more wooded than at present. Paleoclimatic conditions appear to have been as different from historic norms as those seen in several fossil localities in the Western Cape, pointing to greater environmental heterogeneity than has previously been documented for the region in the later Pleistocene.     

Optimal foraging in bumblebees: Calculation of net rate of energy intake and optimal patch choice

To test many predictions of “optimal foraging theory” it is necessary to calculate the rate of net energy intake of a foraging animal. Equations are derived for the calculation of the rate of net energy intake of a foraging bumblebee. The assumptions that form the basis of these energy equations are discussed. As examples, the rates of net energy intake are calculated for Bombus flavifrons workers foraging on neighboring patches of Aconitum columbianum and Delphinium barbeyi. If the bumblebees forage optimally, their net rates of energy intake in the two patches should be equal. The observed rates are consistent with this hypothesis. The application of an optimality approach to pollination biology is briefly discussed.     

A theoretical investigation of the effect of predators on foraging behaviour and energy reserves

Data show that when small birds are exposed to a model of a predator, their body mass may either increase or decrease. Although attempts have been made to explain the data using previous models, these models are based on a constant level of predation and hence are not appropriate for making predictions about the response of a bird to the sight of a predator. We have developed a novel model that includes encounters between a bird and potential predators. We show that, depending on the biology of the predator, optimal body mass may either increase or decrease. The model also makes predictions about the foraging behaviour of the bird after it has seen a predator.     

A comparison of two rotational stocking strategies on the foraging behaviour and herbage intake by grazing sheep

An understanding of the processes involved in grazing behaviour is a prerequisite for the design of efficient grassland management systems. The purpose of managing the grazing process is to identify sward structures that can maximize animal forage daily intake and optimize grazing time. Our aim was to evaluate the effect of different grazing management strategies on foraging behaviour and herbage intake by sheep grazing Italian ryegrass under rotational stocking. The experiment was carried out in 2015 in southern Brazil. The experimental design was a randomized complete block with two grazing management strategies and four replicates. The grazing management treatments were a traditional rotational stocking (RT), with pre- and post-grazing sward heights of 25 and 5 cm, respectively, and a ‘Rotatinuous’ stocking (RN) with pre- and post-grazing sward heights of 18 and 11 cm, respectively. Male sheep with an average live weight of 32 ± 2.3 kg were used. As intended, the pre- and post-grazing sward heights were according to the treatments. The pre-grazing leaf/stem ratio of the Italian ryegrass pasture did not differ between treatments (P > 0.05) (~2.87), but the post-grazing leaf/stem ratio was greater (P < 0.001) in the RN than in the RT treatment (1.59 and 0.76, respectively). The percentage of the non-grazed area was greater (P < 0.01) in post-grazing for RN compared with RT treatment, with an average of 29.7% and 3.49%, respectively. Herbage nutritive value was greater for the RN than for the RT treatment, with greater CP and lower ADF and NDF contents. The total time spent grazing, ruminating and resting did not differ between treatments (P > 0.05), with averages of 439, 167 and 85 min, respectively. The bite rate, feeding stations per min and steps per min by sheep were greater (P < 0.05) in the RN than in the RT treatment. The grazing time per hour and the bite rate were greater (P < 0.05) in the afternoon than in the morning in both treatments. The daily herbage intake by sheep grazing Italian ryegrass was greater (P < 0.05) in the RN than in the RT treatment (843.7 and 707.8 g organic matter/sheep, respectively). Our study supports the idea that even though the grazing time was not affected by the grazing management strategies when the animal behaviour responses drive management targets, such as in ‘Rotatinuous’ stocking, the sheep herbage intake is maximized, and the grazing time is optimized.     

Retinular fine structure in compound eyes of diurnal and nocturnal sphingid moths

Summary Retinular fine structure has been compared in the superposition compound eyes of three sphingid moths, one nocturnal, Cechenena, and two diurnal, Cephonodes and Macroglossum. Cechenena and Cephonodes have tiered retinas with three kinds of retinular cells: two distal, six regular and one basal. The distal retinular cells in Cechenena are special in having a complex partially intracellular rhabdomere not present in Cephonodes. Macroglossum lacks the distal retinular cell. In Cephonodes a unique rhabdom type, formed by the six regular retinular cells in the middle region of the retinula, is divided into three separate longitudinal plates arranged closely parallel to one another. Their constituent microvilli are consequently all nearly unidirectional. The ratio of rhabdom volume to retinular cell volume in the two diurnal sphingids is 10–27%; this is about the same as that (25%) of skipper butterflies, but significantly smaller than in the nocturnal Cechenena (60%). In the diurnal sphingids retinular cell membranes show elongate meandering profiles with septate junctions between adjacent retinular cells. From the comparative fine structure of their eyes the diurnal sphingids and the skippers would appear to be phylogenetically closely related.Supported in part by grants from Ministry of Education Japan (Special Project Research in Animal Behaviors)