BIOLOGY OF THE IMMIGRANT CATTLE EGRET ARDEOLA IBIS IN GUYANA, SOUTH AMERICA

The success of the immigrant Cattle Egret Ardeola ibis in Guyana appears to be due mainly to its breeding biology, combined with man's activities in clearing and draining land for stock and by his selective shioting.
Long.-established heronries are beine taken over bv the Cattle Egret. In Aurmst 1959 during the principal nesting season in the main rains 90% of the 1786 heron nests in the Botanic Gardens Georgetown, belonged to Cattle Egrets, the others to Little Blue Herons Hydranassa caerutea , Black-crowned Night Herons Nycticorac nycticorux , Snowy Egrets Egretta thula , Tricolour Herons Hydranassu tricolor , and Streaked Herons Butorides striatus . In the December rains the same year another 1700 Cattle Egret nests were in use here, but no indigenous herons were nesting. These were, however, joined in January by Black-crowned Night Herons, Snowy Egrets, Little Blue and Tricolour Herons.
In addition to having two peak nesting periods a year, instead of one as the indigenous herons generally have, the Cattle Egrets react faster to the onset of the rains and obtain the best nest-sites, and they are helped by the tendency of the indigenous herons to display more aggressively against their own species than against the intruder; also now that Cattle Egret numbers are high they can find mates very rapidly. Both indigenous herons and Cattle Egrets have one to three young per brood in Guyana, and the nesting period lasts about two months.
The peak nesting time varies from year to year with the advent of the rains. Nesting appears to be stimulated by, and its success related to, rainfall. A cycle of wet years in Guyana preceded the arrival of the Cattle Egrets in North America in 1952. In Guyana their dispersal inland was restricted, probably by the forest, until 1961.     

Seasonal variation in reproductive measures of tropical Roseate Terns Sterna dougallih previously undescribed breeding patterns in a seabird

Seasonal variation in egg-laying, egg size, hatching success, hatchling mass, fledging success and chick growth of Roseate Terms Sterna dougallii breeding on Aride Island (Seychelles), Indian Ocean, were studied in 1997 and 1998. I investigated to what extent two patterns, common in a range of species, were followed by tropical Roseate Terns: (a) seasonal decrease in clutch size, egg size and breeding success and (b) an increase in breeding success with increasing egg weight. In 1997 (a poor year), the earliest nesting birds laid significantly smaller eggs, and chicks were lighter at hatching than those of peak nesting birds. The mean clutch size, of 1.04 eggs, showed no seasonal variation and no 'b'-eggs hatched. In 1998 (a good year) the earliest nesting birds laid eggs of similar size and their chicks were of similar weight to those of peak nesting birds. Mean clutch size, of 1.25 eggs, increased significantly through the season and about 60% of the 'b'-eggs hatched. In 1997, hatching success was 57% whereas in 1998 it was 80%. In both years, breeding success declined significantly through the season. The fact that the earliest breeding birds laid smaller eggs in a poor year and smaller clutches in a good year is in marked contrast to a range of other species, and to temperate-nesting Roseate Terns. Egg volume explained about half of the variance in hatchling mass in both years, but only 15% of the variation in linear growth rate. Hatching date was the only variable with a significant effect on fledging success. Roseate Terns on Aride seemed to sacrifice egg size and clutch size for earliness of laying. Presumably it is a strategy of older birds to lay as early as possible and may be regarded as a response of tropical Roseate Terns to breeding under relatively poor, and seasonally declining, food conditions.     

The breeding biology of the White-spectacled Bulbul,Pycnonotus xanthopygos,at the northwestern edge of its distribution range

The White-spectacled Bulbul, Pycnonotus xanthopygos, is an abundant and possibly invasive species in Turkey, where it has gradually expanded its distribution and breeding range in both western and southeastern directions. This study focused on its breeding biology, which is still poorly known. The breeding activity extends from February until September. The preferred nesting areas are mainly gardens and maquis groves, where 24 different nesting tree species have been identified. The clutch size is 3.3 ± 0.8 eggs per pair, nesting success 68%, hatching success 94%, fledgling success 95%, and overall breeding success 89%. While nesting success differs significantly between the years, we found no significant differences in hatching, fledging, and overall breeding success between the years studied. Despite favourable climatic conditions in the Mediterranean region, the species makes only one brood per year in a relatively extended breeding season extending over seven months, and has a relatively a high reproduction rate per nest.     

Breeding success of the Eurasian eagle owl (<Emphasis Type="Italic">Bubo bubo</Emphasis>) and rodent population dynamics

The dependence of the breeding success of the eagle owl on the population dynamics of rodents, which are a staple of its diet in Mordovia, was traced. At peak numbers of rodents, large clutches and a high survival rate of fledglings were observed in the breeding pairs; after a year of depression of rodent populations, in the following year, the pairs do not nest at all or their breeding success is reduced to a minimum due to the death of the clutches. In 52% of the cases, the nesting pairs laid three eggs; in 31%, two eggs; in 9%, four eggs; in 4%, one egg or five eggs. The average clutch size was 2.78 ± 0.17. The average number of chicks grown from a successful nest was 2.41 ± 0.27. The nests in Mordovia were located at a distance of 1.1–3.7 km from residential areas.     

Changes in clutch size, brood size and numbers of nesting Squacco Herons Ardeola ralloides over a 32-year period in the Camargue, southern France

Changes in numbers of nesting pairs, clutch size, and brood size of Squacco Herons Ardeola ralloides in the Camargue, southern France were analysed over a 32-year period. The annual numbers of breeding pairs [average 84 pairs pL 30 sd, n = 32 years) exhibited a possible increasing trend, but with considerable variability. This variability is associated with local spring rainfall, which may influence the available surface area of their preferred freshwater habitats. In contrast to population size, clutch size decreased substantially in recent years. Our data do not provide a causal explanation for this change in clutch size, although the reduction temporally corresponds with increasing rice cultivation and with a dramatic increase in the number of Cattle Egrets Bubulcus ibis in the Camargue.     

Breeding biology of the Barn Owl Tyto alba in central Mali

Data were obtained on 178 clutches of African Barn Owls in central Mali from four breeding seasons during 1979–1983. Significantly more clutches were laid in 1979–1980 and significantly fewer in 1980– 1981 than the average for the 4 years and there were significantly more clutches laid in the middle period of the annual breeding season. The egg volume was significantly smaller at the beginning of the breeding season and significantly larger in the middle than the overall mean with eggs of second clutches being larger than those of first clutches. The clutch size was 605 eggs of which 479 hatched. The number of young fledged per successful nest was 319 and was 1 83 for all nesting attempts. The month was the only variable shown to affect significantly the clutch size, eggs hatched and fledging rate, the highest success rates being associated with the middle of the breeding period. The average interval between the hatching of eggs was 2–31 days. Survival rates (47'1%) to fledging were significantly affected by year (1981–1982 being the least) and month (mid-season birds the best). The order of hatching significantly affected age at death or disappearance, the first-hatched birds surviving the longest. The year significantly affected age at fledging, the young from the year in which most clutches were laid leaving the nest at the youngest age and those associated with the year having the least number of clutches remaining in the nest the longest. The month of hatching also affected fledging age, birds at the extremes of the breeding season fledging at older ages. The discussion compares these data with those from elsewhere.     

Greater flamingos Phoenicopterus roseus are partial capital breeders

Capital breeding refers to a strategy in which birds use body stores for egg formation, whereas income breeders obtain all resources for egg formation at breeding sites. Capital breeding should occur more in large‐bodied species because the relative cost of carrying stores for egg formation becomes smaller with increasing body size. Based on a comparison between stable isotopes of carbon and nitrogen in potential prey at wintering sites and eggs, we examined whether greater flamingos use nutrients stored earlier in the year for egg production. Our results suggest that the greater flamingo is a partial income breeder, since prey for egg formation were obtained both in overwintering sites and at the breeding site. This may be because there are selective pressures for nesting females to lay soon after arriving at the breeding site, which may be facilitated by arriving at the breeding site with developed ovarian follicles.     

3. THE GOLDEN ORIOLE (Oriolus oriolus) IN HOLLAND

Even though Sfax salina in Tunisia hosts a significant part of the Mediterranean population of Pied Avocet Recurvirostra avosetta, data on the breeding ecology of this species in this area are lacking. Such data are nonetheless necessary, not only for the understanding of the ecology and dynamics of this species in general, but also for conservation purposes. In this paper we use the results of a one-year monitoring of nests in Sfax salina to provide information on its nesting parameters, in particular nesting phenology, colony size and hatching success. Our results show that Pied Avocets formed dense colonies at the beginning of the nesting season, but colony size decreased as the nesting season advanced. Clutch size varied between 1 and 7 eggs, similar to other Mediterranean and European populations. Hatching success, estimated by the Mayfield method, was relatively low, mainly due to predation by stray dogs. Overall, our work shows that relatively high nest losses occur here. It also highlights the need for management efforts aiming to enhance the reproductive success in this artificial habitat. Creation of suitable and safe nesting sites is proposed.     

Nest positioning by male Daito White‐eyes Zosterops japonicus daitoensis improves with age to reduce nest predation risk

Age‐related improvement in reproductive success is widely observed in birds, and the mechanisms by which productivity is enhanced have received considerable attention. However, little is known about how parental age affects the loss of eggs or nestlings despite the fact that age effects on nesting success are often reported. We examined parental age effects on reproductive success in relation to the avoidance of nest predation in an island subspecies of the Japanese White‐eye, the Daito White‐eye Zosterops japonicus daitoensis. Clutch size and annual number of breeding attempts did not differ between parental age classes. Reproductive success was affected only by male age through an increase in nesting success. Nest failure was attributed only to predation in this species and nest concealment and nest height were important nest characteristics promoting successful fledging. Older males built their nests in more concealed and higher positions than first‐year birds, regardless of vegetation status around the nest. Analysis of individual birds suggested that by shifting the nest to a safer position, male White‐eyes achieved higher nesting success than in the previous year. Of three hypotheses of age‐related improvement in reproductive success considered, our data favoured the hypothesis that as individuals grow older, their breeding competence improves.     

Breeding site fidelity and natal philopatry in the Redshank Tringa totanus

This paper presents the results of a study carried out on breeding Redshank in the Ribble Marshes, Lancashire, England.
Redshank tend to return to the same breeding area year after year. There was no detectable sex bias in return rates. Experienced birds were more site faithful than inexperienced birds, with previously successful birds exhibiting the highest degree of breeding site fidelity. Older, more experienced birds were more successful at hatching eggs than inexperienced birds.
Breeding dispersal was the same both within and between years. Faithful pairs and males nesting with a new mate dispersed significantly shorter distances than females nesting with a new partner. Dispersal distances in female Redshank were affected by breeding success: unsuccessful females, nesting with a new mate, dispersed significantly farther than successful females. A pair's breeding success influenced the following year's mate fidelity. However, other factors such as overwintering survival and date of return may also have influenced mate fidelity.
Redshank were highly faithful to their natal area; a high proportion of birds that survived post-fledging mortality returned to breed in their area of birth. No sex bias in natal dispersal was detected. Approximately 50% of Redshank breed in their first year of life.     

BREEDING OF SACRED IBIS THRESKIORNIS AETHIOPICA AT LAKE SHALA, ETHIOPIA

Data are based on more than 200 h of observation at Ethiopia's Lake Shala from 1966 to 1972. Except for differences in size of bill, there are no useful field characters separating male and female Sacred Ibis. The breeding plumage is described; vivid blood-red colour underneath the wings and the ornamental plumes are especially obvious when nesting commences. Physical and biological features of Lake Shala, Ethiopia, and its nesting islands are described; the species of birds nesting on the Shala islands are given. Ibises nest at Shah from March to August; no nesting has been recorded from September to February during the last months of the ‘big’ rains through the main dry season. Nesting normally begins in the ‘small’ rains (between 14 March-24 April), although instances were recorded as early as 1 March and as late as 20 August. The ibises normally nest once per year, although it is possible that occasionally a second nesting may occur after an unsuccessful first attempt. The ibises at Shala nest in discrete groups; several nesting groups may form on any or all of the islands; the number of groups attempting to nest varied from year to year. Nesting activity begins when males arrive and establish pairing territories, usually in a small tree but sometimes on the ground. When females and other males arrive at the pairing territories, pair formation ensues. At this time males perform forward threat, modified forward threat, pursuit flight, supplanting attack and modified snap displays, while both sexes perform stretch and bow displays. Once established, the pair abandons the pairing territory and moves to the nesting area, usually near but always distinct from the pairing territory, and establishes a nest-site territory. Most members of the nesting group move to the nesting area on the same day. Copulation then takes place, and is followed by collection of nest material, usually by the male. Nests are built close together. The average area of 10 nests measured was 0.09 m². Nests are usually less than 20 cm thick and are made of many small branches and sticks. The average clutch in 34 nests was 2.24 eggs; the average size of 34 eggs was 63.4×43.5 mm. Incubation probably begins when the clutch is complete. Both sexes incubate, and the incubation period probably lasts 28–29 days. The development of the young is described. The young leave the nest-site territory when 14–21 days old. Although they are capable of some flight when 35–40 days old, the young do not leave the colony until they are 44–48 days old. In the colony, both parents care for the young. Usually only one parent at a time is with the young. The parents recognize their own young and are usually recognized by them. The behavioural interactions between young and parents are described. Fledging success in 1968 was 1.06 young per pair. The number of pairs successfully rearing young varied annually from none to 81%, on average over six years (1966–70, 1972) 35%. Predation at the breeding colonies is minimal. The food of one one-month old chick consisted of beetle larvae, lepidopteran larvae and beetles. Feeding areas, although undetermined, must be widespread. Inter-specific competition between Sacred Ibis and other nesting birds at Shala is discussed. Among possible factors stimulating nesting at Shala one, fairly heavy rainfall, seems to be especially important. It is also suggested that especially heavy rain-storms cause ibises to abandon the colonies, and result in poor breeding success.     

FREQUENCY-DEPENDENT FITNESS CONSEQUENCES OF INTRASPECIFIC NEST PARASITISM IN SNOW GEESE

The reproductive efficiency, defined as the number of breeding recruits produced per egg laid; of intraspecific nest parasites; of hosts in parasitized nests; and of unparasitized nesting females, was measured for 14 years for lesser snow geese Anser caerulescens caerulescens nesting near Churchill, Manitoba, Canada. Relative efficiencies were 0.71–0.88, 0.91, and 1.0 for eggs of parasites, hosts, and unparasitized birds, respectively. Differences in the hatching probabilities of the three classes of eggs produced the efficiency differences. Parasitic success was limited by the parasites' failure to place more eggs than expected by chance into nests at the appropriate time relative to host incubation. Host nesting success was lower when more than one parasitic egg was added to the clutch. No differences in gosling survival and breeding recruitment probabilities were detected among any categories of goslings. Thus, hatching parasitic young are at no disadvantage relative to parental young, and there is no support for the hypothesis that increased success of host young at later stages of reproduction might offset negative effects at the egg stage. The hatching efficiency of parasitic eggs declined more rapidly than that of parental eggs as the parasitism rate increased. Efficiencies were similar when 3–4% of the eggs laid per year were parasitic, but relative parasitic efficiency was significantly lower when the parasitism rate was 8–9% or more. Using ancillary information and assumptions about the fecundity, viability, and behavioral flexibility of parasitic and parental females, we conclude that intraspecific nest parasitism could compete with nesting as a reproductive strategy in this population. The conditional use of parasitism by a large component of the population in certain years, however, combined with negative-frequency dependent success, limits the potential spread of a purely parasitic strategy in this population.     

Reproductive performance of the Bald Ibis Geronticus calvus in relation to rainfall and grass-burning

Reproductive performance of the Bald Ibis Geronticus calvus was studied during three consecutive nesting seasons (1978–80) at a single large breeding colony in Natal, South Africa. A focal sample of 22 nest sites (out of a total of 60 in the colony) was inspected at frequent intervals to determine clutch sizes, egg survival, hatching success, nestling mortality and fledgling production in each year. Mean clutch size was slightly lower in 1980, compared to the two previous years, although survival and hatching success of eggs were similar in all three years. However, nestling mortality was significantly higher in 1980, largely as a result of parental nest desertion and nestling starvation, so that the proportion of breeding pairs producing at least one fledged offspring, and the average brood size at fledging, were lower compared to 1978 and 1979.
The 1978 and 1979breeding seasons were both preceded by 12 months of above average rainfall, and Bald Ibises fed extensively on small invertebrates obtained in burnt grassland in the study area during the nesting period. By contrast, the onset of nesting in 1980 followed 11 consecutive months of sub-normal rainfall. As a result of the drought, and the consequent lack of controlled burning by landowners in the study area in 1980, very little burnt grassland was available during the Bald Ibis' breeding season, and the birds fed primarily in cultivated pastures during the customary burning period. The lower reproductive performance of Bald Ibises at the study site in 1980 is discussed in relation to the possible effects of drought on the abundance of invertebrate prey and the extent of controlled burning in the species' grassland biotope.     

Are salinas a suitable alternative breeding habitat for Little Terns Sterna albifrons?

This paper describes the breeding population, breeding habitats and reproductive variables of Little Terns Sterna albifrons in natural (sandy beaches) and alternative (salinas) habitats. Studies of nesting success conducted between 1998 and 2002 in these two types of habitat were combined with a literature review of census work from the past 30 years in order to assess whether salinas are suitable alternative breeding habitat for Little Terns. Most of the Portuguese Little Tern population now breeds in salinas. Census data from the last 30 years show that this is a recent breeding behaviour, because until the 1990s most colonies were located on sandy beaches. Destruction and disturbance of the natural habitat has caused this habitat shift. Despite this shift, the Portuguese Little Tern breeding population did not decline during this period and no significant differences were found in nesting success between natural and alternative habitats. This might indicate that salinas are a suitable alternative breeding habitat for Little Terns, but differences in laying period, clutch size and egg size were recorded between birds nesting on sandy beaches and in salinas in the same area. Birds nested earlier on sandy beaches and laid larger clutches and eggs than in salinas. These data suggest that, when both habitats are available, older and/or higher quality birds prefer sandy beaches for breeding, presumably trying to re-nest in salinas when first breeding attempts failed. We discuss conservation priorities and management actions for both habitats.     

Sexually dimorphic eggs, nestling growth and sibling competition in American Kestrels Falco sparverius

1. American Kestrel ( Falco sparverius ) nestlings are sexually dimorphic, with daughters larger than sons. The larger daughters have an advantage during sibling competition for food in excess of their higher per capita food requirements, and we predicted that parents would reduce this competitive disparity by differentially enhancing the growth of sons, specifically by laying them in larger eggs.
2. In a captive breeding population, eggs producing sons were significantly larger than eggs producing daughters; laying order effects were controlled.
3. The influence of sibling egg size ratios on post-natal size relationships persisted through the nesting period, providing parents with a tool to manipulate size-related phenomena in their offspring.     

Can redistribution of breeding colonies on a landscape mitigate changing predation danger?

The reproductive success of colonially breeding species depends in part upon a trade‐off between the benefit of a dilution effect against nestling predation within larger colonies and colony conspicuousness. However, there may be no net survivorship benefit of dilution if smaller colonies are sufficiently inconspicuous. This raises the question about how the size distribution of breeding colonies on a landscape might change as the predation danger for nestlings changes. In southwest British Columbia, Canada, bald eagle Haliaeetus leucocephalus populations have increased exponentially at ～5% per year in recent decades and prey upon nestlings of colonial breeding great blue herons Ardea herodias faninni. Motivated by field data on reproductive success in relation to colony size, modeling is used to ask under which circumstances trading off a dilution benefit against colony conspicuousness can improve population reproductive success. That is, which colonial nesting distribution, dispersed and cryptic versus clumped and conspicuous, best mitigates predation danger on nestlings? When predators are territorial, the modeling predicts a dispersed nesting strategy as attack rate increases, but not as predator numbers increase. When predators are non‐territorial, the modeling predicts a dispersed nesting strategy as predator numbers and/or attack rates increase. When predators are both territorial and non‐territorial, colonial nesting within a predator's territory improves reproductive success when attack rates are low. This suggests nesting in association with territorial predators may offer decreased levels of predation when compared with nesting amongst non‐territorial predators. Thus a change in the colony size distribution of colonially breeding species might be anticipated on a landscape experiencing a change in predation danger.     

Variation in male‐built nest volume with nesting‐support quality,colony, and egg production in whiskered terns

Nest building can represent an energetically costly activity for a variety of animal taxa. Besides, the determinants of within‐species variation in the design of nests, notably with respect to natural and sexual selection, are still insufficiently documented. Based on an observational study, we examined the influence of nesting conditions (nesting‐support quality, colony, laying date, and year) on male‐built nest volume and also its potential role as a postmating sexually selected display in the whiskered tern Chlidonias hybrida. This tern species is a monogamous colonial bird with obligate biparental care breeding on aquatic vegetation. Hence, large nesting platforms are expected to be a selective advantage because they would better withstand adverse environmental conditions and provide a secure structure for eggs. Nest size may also serve as a postmating sexual trait, and variation in egg production would be positively associated with nest size. We found that nest volume was adjusted to different environmental cues. A positive relationship was found between nest volume and nesting‐support quality, indicating that the leaf density of white waterlily is essential for nest stability. Variation in nest volume was not correlated to colony size but varied among colonies and years. Male‐built nest volume was also positively associated with mean egg volume per clutch but not with clutch size. The fitness consequences of building a large nest are yet to be studied, and additional investigations are recommended to better understand whether the activity of males early during breeding season (e.g., nest building and courtship feeding performance) really serves as postmating sexually selected signals.     

Genetic variation,multiple paternity,and measures of reproductive success in the critically endangered hawksbill turtle (Eretmochelys imbricata)

The Yucatán Peninsula in Mexico contains some of the largest breeding groups of the globally distributed and critically endangered hawksbill turtle (Eretmochelys imbricata). An improved understanding of the breeding system of this species and how its genetic variation is structured among nesting areas is required before the threats to its survival can be properly evaluated. Here, we genotype 1195 hatchlings and 41 nesting females at 12 microsatellite loci to assess levels of multiple paternity, genetic variation and whether individual levels of homozygosity are associated with reproductive success. Of the 50 clutches analyzed, only 6% have multiple paternity. The distribution of pairwise relatedness among nesting localities (rookeries) was not random with elevated within‐rookery relatedness, and declining relatedness with geographic distance indicating some natal philopatry. Although there was no strong evidence that particular rookeries had lost allelic variation via drift, younger turtles had significantly lower levels of genetic variation than older turtles, suggesting some loss of genetic variation. At present there is no indication that levels of genetic variation are associated with measures of reproductive success such as clutch size, hatching success, and frequency of infertile eggs.     

Breeding ecology of the Seychelles Black Parrot Coracopsis barklyi

Knowledge of breeding ecology is required for many conservation interventions. The Seychelles Black Parrot Coracopsis barklyi, endemic to the island of Praslin, is vulnerable to extinction. We aimed to improve understanding of C. barklyi breeding ecology to aid conservation planning. We present the results of four years of research, including nesting cavity characteristics and availability, reproductive success, breeding parameters, parental behaviour and reproductive strategy. Thirty-six breeding attempts were studied over the four seasons. Nests were mainly located in Coco de Mer palms Lodoicea maldivica. Deeper cavities with more canopy cover were preferred. There may be a shortage of high-quality nesting cavities in intensive breeding seasons. Average clutch size was 2.2 eggs, incubation period was c. 15 d and egg fertility was 71%. Rats were key nest predators, causing the failure of up to 33% of breeding attempts. The probability of nest success was 53%. At least 57% of fledglings survived their first year. This species breeds cooperatively and practices a highly unusual side-by-side copulation. We discuss the implications of the results in the context of former, ongoing and potential conservation measures for C. barklyi including translocation, invasive species management, nest box provisioning, habitat restoration and further research.     

The Green sea turtle (Chelonia mydas): further observations on breeding in captivity

Mating, nesting and hatching of the Green sea turtle on Mariculture Ltd's farm on Grand Cayman Island, BWI in 1973 were recorded in a previous paper. The results described were the first of their kind, so that events in the farm breeding colony in the following year were awaited with much interest, especially as to hatchability of the eggs. The observations made in 1974, which extended and mainly confirmed those made in 1973, are described below.