Diurnal Variability In The Detection Of Dunlin Calidris alpina

The paper describes aspects of the feeding ecology and behaviour of Lapwings and Golden Plovers in an area southeast of York during the winter of 1984–85. Both species chose winter cereal (autumn-sown winter wheat and barley) in preference to other field types, including grassland. A comparison of winter cereals with Ings (permanent, unimproved alluvial meadows, traditionally farmed and subject to winter flooding) revealed no significant differences in the daily behaviour patterns, feeding rates (pecks per minute) or capture rates of large prey items by Lapwings. Lapwings reached highest numbers in winter cereals 8–¸10 cm high.     

Visual fields and foraging ecology of Blacksmith Lapwings Vanellus armatus

The visual fields of Blacksmith Lapwings Vanellus armatus show the characteristics of visual guided foragers that use precision pecking for prey capture – a binocular field of narrow width and limited vertical extent, with the projection of the bill close to its centre and a large blind area above and behind the head. The topography of the total field, particularly the binocular field, is similar to that of European Golden Plovers Pluvialis apricaria. We suggest that the ‘foot‐trembling’ behaviour associated with foraging in Plovers is not under visual guidance but forces the escape of hidden prey, which is detected when the prey item moves into the binocular field to enable its capture in the bill. Foot‐trembling thus functions to extend the effective foraging area of a bird beyond the limits of its visual field.     

Winter field use and habitat selection by Eurasian Golden Plovers Pluvialis apricaria and Northern Lapwings Vanellus vanellus on arable farmland

Eurasian Golden Plovers Pluvialis apricaria and Northern Lapwings Vanellus vanellus winter in large numbers on Britain's farmland. Previous studies in mixed farming areas showed pronounced preferences for permanent pasture, but increasing numbers of plovers winter in eastern Britain where arable farmland dominates. We show that an area of intensive arable farmland supported significant numbers of both species from October to February in the absence of large areas of pasture. Habitat use varied seasonally as the habitats present changed. Throughout winter, use of cereal crops matched their availability: plovers selected harrow in early winter, sugar beet stubbles in midwinter and other crops in late winter. Pastures were rarely used at any time of the year. Flocks occupied only a fraction of the available fields, concentrating most in large fields with open boundaries and where manure had been applied. Daytime feeding was more likely during cold days after nights with a new moon, short duration of moonlight or low-intensity moonlight. These results show that plovers are currently able to utilize intensive arable farmland but future changes in management, such as cessation of manure applications and reductions in sugar beet cropping, could have detrimental effects.     

Variation in the use of pasture by breeding European Golden Plovers Pluvialis apricaria in relation to prey availability

European Golden Plover Pluvialis apricaria populations have declined across Western Europe. Changes to upland grasslands that are utilized by foraging off-duty birds may have contributed to this decline. The use of enclosed agricultural land by a stable breeding population of Golden Plovers in the South Pennines is described to examine this hypothesis. Fields contained foraging plovers from the end of February to the end of July. Off-duty incubating birds commuted to feed on fields, females during the day and males at night. During the day, adults flew from 6.6 to 7.2 km from the nest to feed, whilst at night birds commuted from 2.4 to 2.7 km. The length of time spent foraging by males was inversely dependent upon daylength. Following hatching, the use of pasture declined, although not as much as in other studies. Arable and improved fields received most use by 15 radiotagged birds, whilst the use of pasture throughout the breeding season was negatively correlated with sward height. Adult diet consisted largely of subsurface prey, particularly larval tipulids, although earthworms were an important component in March and April. The biomass of larval tipulids was strongly correlated with an index of the use of fields. The selection of fields by Golden Plovers is therefore dependent upon prey abundance and availability, and, consequently, is likely to be enhanced by summer grazing and poor drainage. These findings are discussed in the light of recent trends in pastoral management.     

The Effect of Moonlight on Scopoli's Shearwater Calonectris diomedea Colony Attendance Patterns and Nocturnal Foraging: A Test of the Foraging Efficiency Hypothesis

Moonlight is known to affect the nocturnal behaviour and activity rhythms of many organisms. For instance, predators active at night may take advantage from increased visibility afforded by the moon, while prey might regulate their activity patterns to become less detectable. Many species of pelagic seabirds attend their colony only at night, in complete darkness, avoiding approaching their nest sites under moonlight. This behaviour has been most often interpreted as an antipredator adaptation (‘predation avoidance’ hypothesis). However, it may also reflect a lower foraging efficiency during moonlit nights (‘foraging efficiency’ hypothesis). Indeed, moonlight may reduce prey availability because preferred seabird prey is known to occur at higher depths in moonlit nights. Using high‐accuracy behavioural information from data loggers, we investigated the effect of moonlight on colony attendance and at‐sea nocturnal foraging in breeding Scopoli's shearwaters Calonectris diomedea. We found that birds departing for self‐feeding trips around the full moon performed longer trips than those departing around the new moon. On nights when the moon was present only partly, nest burrow entrances took place largely in the moonless portion of the night. Moreover, contrary to predictions from the ‘foraging efficiency’ hypothesis, nocturnal foraging activity increased according to moonlight intensity, suggesting that birds increased their foraging activity when prey became more detectable. This study strengthens the idea that colony attendance behaviour is strictly controlled by moonlight in shearwaters, which is possibly related to the perception of a predation risk.     

Spawning migrations in the monogamous butterflyfish,Chaetodon trifasciatus

Spawning and related behavior of a monogamous butterflyfish,Chaetodon trifasciatus, were observed at Kuroshima Island, Okinawa, Japan. Each heterosexual pair defended a feeding territory in the daytime. Spawning occurred at dusk on the days around full or new moon in the daytime. Spawning occurred at dusk on the days around full or new moon in the vicinity of offshore tidal currents. Spawning migration to such sites occurred in pairs, the feeding territories of which were located in areas of inshore currents. In the evening each pair established a small temporary territory, which they spawned adjacent to and slept within until the next morning. The distribution of sleeping sites as well as tidal current directions may determine the spawning sites of this butterflyfish.     

Behaviour of young Ringed Plovers Charadrius hiaticula and its relationship to growth and survival to reproductive age

The behaviour, growth and survival of Ringed Plover chicks were studied in 1974 at Mestersvig, northeast Greenland, and from 1973 to 1976 at Lindisfarne, northeast England. Post-fledging survival, dispersal and recruitment were also investigated for the Lindisfarne population. Growth rates were similar in both study areas, and to those reported from other sites. A diurnal rhythm in feeding activity was more evident in the Arctic than at temperate latitudes, despite continuous daylight in the former area. This was correlated with low nocturnal temperatures and prey availability. It is concluded that food supply did not limit growth or chick survival in either study area. Brooding decreased progressively during the pre-fledging period; the relationship of brooding period to age, environmental conditions and area was investigated. The adaptations of feeding and brooding behaviour for arctic and temperate breeding Ringed Plovers are discussed. Survival from hatching to fledging varied between 40% and 60% in different areas and years, and at least 59% of birds survived from fledging to about one year old. Minimum survival from one to two years old was only 57%. but year-to-year survival of older birds was higher, that of breeding adults being at least so?;. Most birds returned to the breeding area and attempted to breed when one year old. The extent of dispersal from Lindisfarne and the wintering areas of the Lindisfarne-breeding birds are investigated.     

Celestial moderation of tropical seabird behavior

Most animals, including birds, have cyclic life histories and numerous studies generally conducted on captive animals have shown that photoperiod is the main factor influencing this periodicity. Moon cycles can also affect periodic behavior of birds. Few studies have investigated the influence of these environmental cues in natural settings, and particularly in tropical areas where the change in photoperiod is slight and some bird species keep cyclic behaviors. Using miniaturized light sensors, we simultaneously investigated under natural conditions the influence of photoperiod and moon phases on the migration dates and at-sea activity of a tropical seabird species, the Barau's petrel, throughout its annual cycle. Firstly, we found that birds consistently started their pre- and post-breeding migrations at precise dates corresponding in both cases to a day-duration of 12.5 hours, suggesting a strong influence of the photoperiod in the regulation of migration behavior. We also found that mean population arrival dates to the colony changed from year to year and they were influenced by moon phases. Returns at their colonies occurred around the last full moon of the austral winter, suggesting that moon cycle is used by birds to synchronize their arrival. Secondly, variations of day-time activity were sinusoidal and correlated to seasonal changes of daylength. We thus hypothesize that the photoperiod could directly affect the behavior of the birds at sea. Night-time at-sea activity exhibited a clear cycle of 29.2 days, suggesting that nocturnal foraging was highly regulated by moon phase, particularly during the non-breeding season. To our knowledge, this is the first study to document a mixed regulation of the behavior of a wild bird by photoperiod and moon phases throughout its annual cycle.     

ASPECTS OF THE BREEDING BIOLOGY OF THE FIERYNECKED NIGHTJAR

Jackson H. D. 1985. Aspects of the breeding biology of the Fierynecked Nightjar. Ostrich 56: 263–276.

A marked population of nightjars in Zimbabwe was studied intensively for four breeding seasons. This paper covers certain aspects of the breeding biology of the Fierynecked Nightjar Caprimulgus pectoralis. The male shows strong site fidelity during the breeding season (September to December), singing, feeding and breeding within an area of about 5,8 ha. There is some evidence of site defence by the male. The female shows strong mate fidelity, resulting in a pair bond for life. Egg laying starts with full moon in September and is further stimulated by the next two full moon periods. The eggs are laid directly on dense leaf litter at a site overhung by foliage. The normal clutch is two eggs (12S % are one egg) laid on successive days during the afternoon. Incubation starts with the first egg and is by the male at night and the female by day. The incubation period is 18 days. The birds respond to undue disturbance by deserting the eggs and laying a replacement clutch. The chicks usually hatch on successive afternoons; they are mobile on the first day and will move to a parent if called. Both parents feed and brood the young during twilight and moonlight; the male broods them on dark nights and the female does so by day. The species is double-brooded when time permits, the female laying again once the first brood has reached independence; she may even lay a third clutch if the second one comes to grief. There is no evidence of adults transporting eggs or young.     

The energetic implications of precocial development for three shorebird species breeding in a warm environment

We investigated the effect of body size, parental behaviour and timing of breeding on the chick growth and energetics of three precocial shorebird (Charadrii) species in a warm sub-tropical environment: the Kittlitz's Plover Charadrius pecuarius , Blacksmith Lapwing Vanellus armatus , and Crowned Lapwing V. coronatus . The three species exhibited slower growth, longer pre-fledging periods and lower energy expenditure during the pre-fledging period than Arctic and temperate zone relatives of similar size. It was found that relative growth rates of charadriid and scolopacid species increased by 1.77% per degree latitude. Blacksmith and Crowned Lapwings are of similar size (i.e. about 160 g), about four times the mass of Kittlitz's Plovers. Taking body mass into account, the resting metabolic rate and daily energy expenditure of the three study species were similar. The relative amounts of energy needed for chicks to grow from a hatchling to a fledgling were low for the three species studied. The relative amount of energy needed to complete growth in charadriid and scolopacid species increased by 2.54% per degree latitude. Because Crowned Lapwings exhibited faster growth than Blacksmith Lapwings but similar daily energy expenditure we suggest that Blacksmith Lapwings compensate for greater energy expenditure in colder weather by growing more slowly. We suggest that small growth rate coefficient, low resting metabolic rate and low daily energy expenditure are adaptations to lower food availability and milder ambient temperatures in sub-tropical breeding habitats compared to temperate or Arctic regions.     

OBSERVATIONS ON THE BREEDING BIOLOGY OF THE FISCAL SHRIKE

Ward, D. 1989. Behaviour associated with breeding of Crowned, Blackwinged and Lesser Blackwinged Plovers. Ostrich 60: 141–150.

The behaviour of Crowned Plovers Vanellus coronatus, Blackwinged Plovers V. melanopterus and Lesser Blackwinged Plovers V. lugubris in mate and territory acquisition and defence was documented and related to the habitats these birds occupy. The open habitat occupied by vanelline plovers makes them particularly vulnerable to predation and as a result, they have a highly-developed ability to detect potential predators and have developed a number of behavioural strategies to avoid predation. This has resulted in these birds having a higher reproductive success than that documented for other precocial birds.     

Impacts of “supermoon” events on the physiology of a wild bird

The position of the Moon in relation to the Earth and the Sun gives rise to several predictable cycles, and natural changes in nighttime light intensity are known to cause alterations to physiological processes and behaviors in many animals. The limited research undertaken to date on the physiological responses of animals to the lunar illumination has exclusively focused on the synodic lunar cycle (full moon to full moon, or moon phase) but the moon's orbit—its distance from the Earth—may also be relevant. Every month, the moon moves from apogee, its most distant point from Earth—and then to perigee, its closest point to Earth. Here, we studied wild barnacle geese (Branta leucopsis) to investigate the influence of multiple interacting lunar cycles on the physiology of diurnally active animals. Our study, which uses biologging technology to continually monitor body temperature and heart rate for an entire annual cycle, asks whether there is evidence for a physiological response to natural cycles in lunar brightness in wild birds, particularly “supermoon” phenomena, where perigee coincides with a full moon. There was a three‐way interaction between lunar phase, lunar distance, and cloud cover as predictors of nighttime mean body temperature, such that body temperature was highest on clear nights when the full moon coincided with perigee moon. Our study is the first to report the physiological responses of wild birds to “supermoon” events; the wild geese responded to the combination of two independent lunar cycles, by significantly increasing their body temperature at night. That wild birds respond to natural fluctuations in nighttime ambient light levels support the documented responses of many species to anthropogenic sources of artificial light, that birds seem unable to override. As most biological systems are arguably organized foremost by light, this suggests that any interactions between lunar cycles and local weather conditions could have significant impacts on the energy budgets of birds.     

Nocturnal feeding by Dark-bellied Brent Geese Branta bernicla bernicla

The occurrence and extent of nighttime feeding by Dark-bellied Brent Geese Branta bernicla bernicla on salt marshes adjacent to their tidal roost site were assessed on the north Norfolk coast using position-sensitive radio transmitters. Nine birds were monitored on 143 bird-nights, and feeding was recorded on 87. The frequency of feeding by night varied widely between individuals. The mean proportion of the night spent feeding for one bird was 19.7 ± 8%. Of this, 59% occurred within ± 1.5 h of high tide. Neither the proportion of the whole night spent feeding nor the intensity of feeding around high tide varied with the length of the night or with time after sunset at which high tide occurred. The extent of nocturnal feeding was also independent of the brightness and duration of moonlight. The birds fed for significantly longer on colder nights, and there was a significant positive relationship between the proportion of the night spent feeding and maximum temperature the preceding day. Analysis of daytime activity budgets on inland pastures showed that the proportion of time engaged in nonfeeding, energy-expensive activities was also positively related to maximum daytime temperature. Increases in the proportion of the night spent feeding after warmer days were achieved by increasing the period of time spent feeding on either side of the high tide, not by increasing feeding intensity at high tide. It is concluded that the geese compensate for increased energy expenditure during the day by increasing the extent to which they feed at night. Implications of increasing energy expenditure as a result of disturbance during the day are discussed.     

BREEDING BIOLOGY OF THE MANX SHEARWATER PUFFINUS PUFFINUS

Studies on the breeding biology of Puffinus puffinus were carried out in 1963 and 1964 at the large colony on Skokholm, Wales. During the six weeks before laying the birds spent up to a quarter of the days in the burrows, but the ten days immediately prior to laying were normally spent at sea. There is a prolonged laying period, with a marked peak in the first half of May. Details are given of a second egg being laid when the first was deserted immediately after being laid. The male took the first incubation spell. The incubation spells ranged from one to 26 days and averaged six. The incubation period was about 51 days. The frequency of visits to land by breeding birds, unlike those by non-breeders, was not affected by the moon. On hatching, the chicks grew rapidly and reached maximum weights of between 505 and 755 gm. sometime between 39 and 61 days. There was a variable desertion period, usually eight or nine days, before the chicks left the island about 70 days after hatching. During the feeding period the chick received about two feeds every three days. There is evidence that adults visited the chicks more frequently than this. There was no correlation between growth of the chicks, their feeding rates or fledging weights and the time of laying. There was a high survival (about 95 %) of chicks during the fledging period but some eggs were lost in disputes for burrows. Nine pairs in 1964 were unable to raise two young simultaneously. Parents altered their feeding rhythms to try to feed two young but did not themselves lose weight. It is suggested that the critical factor in the production of young is the availability of food for the young immediately after they leave the colonies.     

Differential moult patterns in relation to antipredator behaviour during incubation in four tundra plovers

Golden plovers and Grey Plovers Pluvialis spp. all have very distinct breeding plumage rich in contrast, with a conspicuous black belly and breast bordered by a bright white fringe. Eurasian Golden Plovers are known partly to replace their breeding plumage with striped yellow feathers during incubation, different from both breeding and non-breeding plumages. In this study a similar partial breeding moult was observed in Pacific Golden Plovers and American Golden Plovers caught on the nest or collected during incubation, although the feathers did not differ clearly from those of non-breeders. This moult starts during incubation and precedes the post-breeding moult into non-breeding plumage. Because the lighter feathers reduce the contrast between the black belly and the white flanks, we suggest that during incubation the plumage characteristic that plays an important role in mate choice is no longer important; at this stage it is better for the bird to be inconspicuous. Additional information on museum skins of golden plovers and of Grey Plovers indicated that only the three golden plovers undergo this partial moult, but that Grey Plovers in general retain full breeding plumage throughout incubation. The three golden plovers also resemble each other in their generally very passive nest defence strategies. In contrast, the larger Grey Plovers actively chase and attack aerial and ground predators. Thus, a reduced conspicuousness of the body plumage during incubation is likely to benefit the golden plovers more than the Grey Plover. We suggest that nest defence behaviour, plumage characteristics and perhaps size have co-evolved as a response to different selection pressures in golden plovers and Grey Plover, but alternative hypotheses are also discussed.     

Lunar influence on prey availability,diet shifts and niche overlap between Engraulidae larvae in tropical mangrove creeks

The influence of the lunar cycle on prey availability, diet shifts and overlap between larval Anchovia clupeoides and Cetengraulis edentulus was evaluated in mangrove creeks of the Goiana Estuary. Copepod eggs were highly abundant in the first and last quarter, at the full moon and zoea of Ucides cordatus (Ocypodidae) in the new moon. The Engraulidae larvae fed on microcrustaceans, algae and early planktonic stages of benthic organisms. The relative importance of prey varied according to prey availability in all moon phases. Larval diets were more even in the full and new moons, when the relative importance of calanoid copepods and zoeae of U. cordatus as food items increased (index of relative importance, >80% I_RI). Mangrove creeks were very important feeding grounds for engraulid larvae during spring tides. Larval diets were more diverse in the first and last‐quarter moon and included protozoeae of Caridean shrimp, larvae of Anomalocardia brasiliana (Veneridae), Isopoda, Gastropoda, ephippium of Daphnia sp. and nauplii of Cirripedia, Harpacticoidia and cyclopoid Copepoda. The last five items were not found in the creeks, suggesting feeding in the main channel. During neap tides, mangrove creeks were probably also used as refugia. These larvae are opportunistic and feed on highly available prey and both species feed on the same items, leading to high dietary overlap in all moon phases. The lunar cycle, which is related to the spring‐neap tidal cycle, was the major driver of quantitative and qualitative changes in feeding of engraulid larvae on a short time scale.     

Negative impact of wind energy development on a breeding shorebird assessed with a BACI study design

Previous studies have shown negative associations between wind energy development and breeding birds, including species of conservation concern. However, the magnitude and causes of such associations remain uncertain, pending detailed ‘before‐after‐control‐intervention’ (BACI) studies. We conducted one of the most detailed such studies to date, assessing the impacts of terrestrial wind energy development on the European Golden Plover Pluvialis apricaria, a species with enhanced protection under European environmental law. Disturbance activity during construction had no significant effect on Golden Plover breeding abundance or distribution. In contrast, once turbines were erected, Golden Plover abundance was significantly reduced within the wind farm (?79%) relative to the baseline, with no comparable changes in buffer or control areas. Golden Plovers were significantly displaced by up to 400 m from turbines during operation. Hatching and fledging success were not affected by proximity to turbine locations either during construction or operation. The marked decline in abundance within the wind farm during operation but not construction, together with the lack of evidence for changes in breeding success or habitat, strongly suggests the displacement of breeding adults through behavioural avoidance of turbines, rather than a response to disturbance alone. It is of critical importance that wind farms are appropriately sited to prevent negative wildlife impacts. We demonstrate the importance of detailed BACI designs for quantifying the impacts on birds, and recommend wider application of such studies to improve the evidence base surrounding wind farm impacts on birds.     

Lunar influences in the diet habits and selectivity of larval clingfish Gobiesox marmoratus

The influence of the lunar cycle on the feeding habits, feeding success and selectivity was tested in situ on larval stages of the clingfish Gobiesox marmoratus (Gobiesocidae). Gobiesox marmoratus larvae and their prey were collected in the water column during the lunar cycle in austral spring 2015 and 2016, in shallow waters (<30 m depth) of El Quisco, central Chile. Feeding incidence was high (80–100%) throughout the moon cycle. The diet was composed of 32 prey items, being gastropod larvae and invertebrate eggs the most important. Prey composition varied among lunar phases with the lowest prey richness during new moon. During 2016, G. marmoratus larvae ingested the lowest number, but the largest prey during new moon. In spring 2015, there were no significant changes in the abundance of each prey taxon in the water column among moon phases, but in spring 2016 there were significant differences in the abundance of cypris and copepod nauplii, particularly between full and new moon. Feeding selectivity index showed that gastropod larvae were positively selected throughout the lunar cycle during spring 2015. In spring 2016, invertebrate eggs were selectively ingested in full moon and third quarter, but at a new moon G. marmoratus larvae selected gastropod larvae. Mean temperature of the water column and its vertical gradient and nocturnal cloud cover influences the feeding success of larval G. marmoratus. Lunar illumination favoured only an increase of richness of prey items. Therefore, nocturnal cloud cover precludes a greater influence of lunar illumination in the larval trophic ecology of this crypto-benthic fish.     

Aktivit?tsrhythmus von Brandg?nsen (Tadorna tadorna) und Watv?geln (Charadrii) an der Nordseeküste

Shelducks, Oystercatchers, Grey Plovers, Knots, Dunlins, Bar-tailed Godwits, Curlews, Spotted Redshanks, Redshanks, and Greenshanks showed a tidal rhythm of time-activity budgets on the Wadden Sea coast during the non-breeding season. Ringed Plovers and Ruffs did not react to the tidal rhythm. Night-time feeding could be observed in most species. Differences in the time-activity budgets of the species were related to body mass and size of food particles. Inter-site comparisons revealed that technical constructions for coastal protection influenced the activity rhythms of the birds.     

State uncertainty models and mark‐resight models for understanding non‐breeding site use by Piping Plovers

Conservation of beach‐nesting medium‐distance migrants has focused on breeding areas because protection of nests is more tractable than protection of non‐breeding habitat. As breeding ground management has encountered diminishing returns, interest in understanding threats in non‐breeding areas has increased. However, robust estimates of non‐breeding demographic rates and abundance are generally lacking, hindering the study of limiting factors. Estimating such rates is made more difficult by complex population dynamics at non‐breeding sites. In South Carolina, endangered Piping Plovers Charadrius melodus start arriving in July and some depart prior to December (the autumn‐only population) while others remain through at least March (the wintering population). State uncertainty capture‐mark‐recapture models provide a means for estimating vital rates for such co‐occurring populations. We estimated the proportion of the population entering the study area per survey (entry probability) and proportion remaining per survey (persistence rate) for both populations during autumn, and abundance of the wintering population, at four sites in South Carolina in 2006/7 and 2007/8, taking advantage of birds previously colour‐ringed on the breeding grounds. We made fairly precise estimates of entry and persistence rates with small sample sizes. Cumulative entry probability was ~50% by the end of July and reached 95% for both populations by October. Estimated stopover duration for birds in the autumn‐only population was 35 days in year 1 and 42 days in year 2. We estimated a wintering super‐population size of 71 ± 16 se birds in the first year and 75 ± 16 in the second. If ringing programmes on the breeding grounds continue, standardized resighting surveys in the non‐breeding period and mark‐recapture models can provide robust estimates of entry and persistence rates and abundance. Habitat protection intended to benefit non‐breeding Piping Plovers at our coastal sites should be in effect by late summer, as many birds are resident from July to the end of winter.