Restoration and management for cliff‐nesting birds in Mediterranean mining sites: the Sand Martin case study

Habitat enhancement for birds is frequently implemented during mine site restoration. Cliff‐nesting birds often colonize anthropogenic environments such as mining areas (aggregate sites and quarries for aggregate and cement production). Mining activity can compromise breeding success, causing cliff‐nesting birds to depend on the management and restoration of mining areas. The objective of our study is to assess the importance of mine site habitats for Sand Martin conservation and reconcile mining activity with breeding success in Mediterranean environments. We studied Sand Martin breeding habitat preferences in mining areas at three spatial scales. At the mining site scale, we compared 10 mining sites with Sand Martin burrows with 19 mining sites without burrows. At the colony scale (vertical structures with colonies), we evaluated the relationships between the number of breeding pairs, number of burrows, and colony characteristics within 30 distinct Sand Martin colonies. At the burrow scale, we compared the characteristics of the available vertical structure with the areas used by Sand Martins. At the mining site scale, Sand Martins preferred more surface of water bodies, shorter distances to flowing water, older sites, and mining sites which produce aggregates instead of cement. At the colony scale, Sand Martins preferred southwest orientations and stockpiles to vertical extraction faces. At the burrow scale, birds preferred the most vertical areas of the face. Our results support the need for effective habitat restoration and improved management for more effective Sand Martin conservation within mining areas. Simple interventions can enhance habitat quality and conservation of cliff‐nesting birds.     

Seasonal roosts of Red-lored Amazons in Ecuador provide information about population size and structure

ABSTRACT.   Data from roosts of Amazona parrots may be useful in creating demographic models, because these birds exhibit high roost fidelity and pairs are conspicuous in flight. However, few investigators have attempted to track changes in the number of pairs using such roosts. We studied Red-lored Amazons ( Amazona autumnalis ) at a communal roost in southwest Ecuador over a 1-yr period to understand better their population structure. Population size was estimated at 214 individuals. Counts revealed seasonal variation in numbers, but the occurrence of pairs and singles was seldom correlated. The number of paired individuals using the roost was lower during the breeding period. In contrast, the number of single birds at the roost nearly doubled during the breeding period. Overall, our data suggest that parental responsibilities during the nesting period explain fluctuations in the number of birds at the roost, and such fluctuations can be used to estimate the reproductive portion of the population. Protection of the small mangrove islands where the parrots roost would likely benefit a population that occupies a much larger area and would, at the same time, provide a useful tool for demographic studies of this poorly known neotropical parrot.     

THE ANNUAL RE-OCCUPATION OF BREEDING SITES BY THE FULMAR

The Fulmar has a long period at the breeding colony prior to egg-laying. The pattern of annual occupation and build-up in numbers has been examined in detail at Marsden, Co. Durham, at a colony in which over 100 eggs are laid annually (Order 3 of Fisher's classification). The re-occupation of the cliff starts in early November with an occasional visit by one or two birds. The main period of activity at the cliff is during the morning and, as the numbers build up, the diurnal period of occupation increases. By mid-December the first birds to arrive in the colony do so before dawn and the last to leave remain well after dark until near midnight. Almost throughout the pre-egg stage, the colony is deserted each night and re-occupied the next day and birds only stay regularly overnight just before egg-laying. A similar pattern of occupation occurs after breeding but in the reverse order. The numbers of birds at the colony in January and February exceed the breeding population and include many non-breeders. The non-breeders progressively decline in numbers until May when only the breeding birds remain with a few non-breeding birds. The daily variation in the numbers of birds at the cliff is influenced by the wind speed. In general, the birds leave the colony under freshening conditions and the number present at the colony can be interpreted in terms of the wind conditions over the last three days. It is suggested that the synchronised departures are primarily feeding trips, the birds using the strong winds to reach feeding areas, except that the departure just before egg-laying is linked to egg development and synchronised laying in the colony. Competition between Fulmars and Kittiwakes for nesting sites usually results in the Kittiwakes gaining the site. This is achieved by the Kittiwakes taking over the Fulmar sites during one of the latter's departures.     

Communal roosting,thermoregulatory benefits and breeding group size predictability in cooperatively breeding sociable weavers

Extreme temperatures impose energy costs on endotherms through thermoregulation and different adaptations help individuals to cope with these conditions. In social species, communal roosting and huddling are thought to decrease the energetic requirement of thermoregulation under low temperatures. This is likely to represent an important mechanism by which individuals save energy during the coldest parts of the year and hence to represent a non‐breeding benefit of sociality. Here, we investigate the potential thermoregulatory benefits of group living in roosting groups of sociable weavers Philetairus socius, a colonial cooperatively breeding passerine that builds communally a massive nest structure with several independent chambers wherein individuals breed and roost throughout the year. To investigate the benefits of sociality during the non‐breeding season, we studied the thermal environment during roosting in relation to group size. In addition, to understand the link between non‐breeding and breeding sociality in this species we studied group size stability between the pre‐breeding and breeding periods. As expected, we found that the nest chamber's night‐time temperature is strongly related to the number of birds roosting together, especially during cold nights. Specifically, birds in larger groups spent less time below the critical thermal minimum temperature (i.e. the temperature below which energy expenditure increases substantially). They were less exposed to external temperature variations. We also found a positive relationship between the number of birds roosting during winter and the breeding group size, indicating breeding group size predictability. In cooperative breeders such as the sociable weaver, the costs and benefits of sociality are usually studied during the breeding period. This study shows that a better understanding of non‐breeding benefits of group membership and group dynamics between the non‐breeding and breeding periods are necessary for a comprehensive understanding of the benefits of sociality.     

COMMUNAL ROOSTING BEHAVIOUR OF THE AUSTRALASIAN HARRIER CIRCUS APPROXIMANS IN NEW ZEALAND

The Australasian Harrier Circus upproximans habitually roosts communally in New Zealand but not in Australia. As many as 100 birds can occupy a roost in a small area of swamp. They start to assemble about one hour before dark and communal aerial displays in the vicinity of the roost are an integral part of the roosting behaviour. Roosts may be occupied all the year round, non-breeding birds continuing to roost communally throughout the summer. In New Zealand the habit is neither connected with migration nor is it an anti-predator device. It is considered that the abundance of food caused by the spread of introduced mammals and the large increase of habitat created by European settlement has built the harrier population up to such a size that the number required to elicit the response of communal roosting occurs at all times of year in most districts. Numbers within individual roosts fluctuate throughout the year and it is suggested that communal displays before roosting may provide the necessary feed-back for the initiation of population adjustments.     

OBSERVATIONS ON THE COMMUNAL ROOSTING OF AFRICAN BLACK OYSTERCATCHERS

Hockey P. A. R. 1985. Observations on the communal roosting of African Black Oystercatchers. Ostrich 56: 52–57.

There are currently three main hypotheses for the adaptive significance of avian communal roosts: physiological advantages, predator avoidance and information centres. The African Black Oystercatcher Haematopus moquini is territorial throughout the year and forms communal roosts during the nonbreeding season, but does not breed communally. Roosts generally are small, and site fidelity is high. Roosts are normally sited on a rocky promontory with adjacent offshore rocks (west coast), or in flat areas with extensive all-round visibility. The main predators of African Black Oystercatchers are nocturnal terrestrial mammals and nocturnal communal roosts are larger and more tightly packed than daytime roosts. Breeding birds do not roost communally during the breeding season and at this time of year mortality due to mammal predators at a study site in Saldanha Bay was greatest (X² = 9.46; p<0.01). It appears therefore that predator avoidance is an important adaptive feature of communal roosting in this species.     

PHOTORESPONSES OF THE WOODPIGEON COLUMBA PALUMBUS IN RELATION TO THE BREEDING SEASON

• 1 The gametogenetic response following photo-stimulation was tested in Woodpigeons under controlled laboratory and aviary conditions. Subjects were wild-caught adult or first-winter birds or were hand-reared from wild-taken nestlings. The natural seasonal gonad cycle is described and experiments were related to this cycle.
• 2 Gonad recrudescence could be initiated in adults with regressed organs by exposure to summer photoperiods, while the normal vernal recrudescence could be prevented if birds were artificially kept on short (winter) daylengths. Thus Woodpigeons could be stimulated towards reproductive condition at seasons when the gonads of wild-living birds remained inactive, viz. late September to mid-February.
• 3 Juvenile males were unresponsive to extra photo-stimulation until they were about six months old. This meant that some spermatogonial division could be procured in a few individuals in late February, but juveniles were not very noticeably affected by an extended photoperiod until mid-late March. This is the time when juveniles first begin gametogenetic development in the wild, though spermatogenesis normally accelerates in April after which time the rate of gonad development is constant. First-winter females were judged to resemble males in their photoperiodic responses but the sample examined was small.
• 4 The results confirm that the natural gonad cycle in the Woodpigeon is controlled (via gonadotrophic hormones) by seasonal changes in daylength. More important is the finding that the species appears to possess no post-nuptial refractory period. Thus, at the end of the breeding season in September birds kept on summer daylengths were maintained in full reproductive condition until December. The testes of controls placed on winter daylengths regressed within one month, but they were immediately responsive to a summer daylength which stimulated spermatogenesis within a month.
• 5 The discussion argues that the avian refractory period is not a necessary period for gonad rehabihtation and reorganization and that it has not evolved as a fixed period serving to time the breeding season; views which until now have been current. Instead, it functions only as a safety mechanism, preventing unseasonal reproduction in those species for which natural selection favours seasonal breeding. In such species the evolution of sensitivity to a particular daylength could result in breeding taking place at the wrong time, for example spring responding species might also respond in the autumn. Because natural selection permits Woodpigeons to remain in breeding condition from March until September, without disadvantage, a direct response to natural daylength can regulate the cycle, and there is no need for a period of pituitary refractoriness. It is not known if some pigeons have lost the refractory period or whether they never possessed one.

     

Linking anti-predator behaviour and habitat quality: group effect in nest defence of a passerine bird

In habitats where the density of breeding individuals is higher, breeding success has been shown to increase with the number of close conspecific and heterospecific neighbours. However, the mechanisms linking habitat quality, group size of prey individuals and offspring defence are poorly known. In this field study, we examined the relationships between habitat quality and parental nest defence behaviour in the pied flycatcher (Ficedula hypoleuca). We found that mobbing is more intense in unmanaged forests where birds breed in more dense and diverse communities than in heavily managed young forests where heterospecific densities are lower. We also found that the mobbing activities of pied flycatchers breeding in unmanaged mature boreal forests attracted more neighbouring prey individuals than in nearby managed forests. This study shows that habitat quality-mediated effects might be responsible for the decreased group size of mobbing birds in managed forests, which may lead to less effective communal defence.     

Cladistic analysis of cassiduloid echinoids: trying to see the phylogeny for the trees

Using DNA fingerprinting we estimated the reproductive success of 49 adult birds belonging to 16 breeding groups of the sexually monomorphic brown skua (Calharacta lonnbergi) from the Chatham Islands (New Zealand). This population has a variable mating system, breeding in both monogamous and polyandrous groups. The parentage of 45 chicks produced over three breeding seasons was unequivocally determined using the multilocus probes 33.15 and 33.6. We found no evidence of either extra-pair or extra-group fertilization and there was no evidence to suggest egg dumping by females in any breeding group. Consequently, in the case of pairs, parentage of all chicks was assigned to the resident adult birds. In addition, band sharing analysis indicated that members of communal groups were not close relatives. In the 10 communally breeding groups examined, multiple paternity within a clutch was recorded on two of the 12 occasions in which two chicks were reared. Analysis of the parentage of offspring belonging to different groups, from different years, demonstrated that the number of chicks produced by some adult males varied considerably between seasons. In contrast, the reproductive success of other individuals was constant; for example, one male produced two chicks in each of the three seasons it was studied, while other males in communal groups did not produce any chicks during the course of this study. Fitness is a lifetime parameter, and any assessment of it requires studies over at least the average lifetime of an individual. The findings presented in this study suggest that, for brown skuas, there are significant differences in the reproductive success of some adult males in different breeding seasons. These results indicate that estimates of reproductive fitness based on only a single breeding season's data can be seriously inaccurate. Should temporal changes in paternity (and/or maternity) be shown to be common phenomena in other species, then such results would have major implications for the interpretation of parentage studies.     

FACTORS AFFECTING THE AGE OF FIRST BREEDING OF THE KITTIWAKE RISSA TRIDACTYLA

At a Kittiwake colony in Northumberland, 80% of those birds which returned to their natal colony to breed were males and these supplied 52% of all male recruits. More females breed away from their natal colony than males. There was no differences in the proportions of young fledged from sites in the centre or at the edge of the colony, or by parents of different experience, which returned to breed. Kittiwakes breed for the first time at ages from 3 to 8 years, but most at 4 or S years old. Males arrive back at the colony at an earlier age than females and breed for the first time one year earlier. Males obtaining sites at the centre of the colony first breed at an earlier age than those at the edges. Neither the age nor the area of first breeding appear to be transmitted from parent to offspring. Males breeding first aged 4 years or younger produced more young than those which first bred aged 5 years or older, despite their partners laying smaller clutches. This difference was most marked among those males recruited to sites in the centre of the colony. The advantage of this earlier breeding is counteracted by a lower survival rate among those males which start to breed at the younger ages. In all breeding Kittiwakes, annual reproductive output increases with experience while annual survival rates decrease. Once they had started to breed, many birds failed to breed in one subsequent season. Nearly 60% of these cases of intermittent breeding occurred in the year following first breeding. Intermittent breeding was most frequent among young birds and among females. It is suggested that each breeding involves a cost to the individual in terms of reduced survival, and that deferred and intermittent breeding are means of guarding survival. A model is proposed whereby the age at which a bird starts to breed, the nesting site which it obtains, and its subsequent breeding strategy result in each individual producing an optimal number of reproducing offspring in its lifetime, relative to its quality.     

FOOD AS A FACTOR CONTROLLING THE BREEDING OF PUFFINVS LHERMINIERI

A study of Puffinus Iherminieri on Plaza Island, Galápagos, showed that, although eggs were laid in all months, there were marked peaks and troughs of laying. Intervals between successive layings varied with the success or failure of the first egg, an average of nine months for successful pairs, eight months for pairs which lost a young, and 6–5 months for those which failed to hatch an egg. However the breeding success did not influence the time between the end of one breeding attempt and the next laying. This suggests that birds were breeding as quickly as possible. The critical factor preventing more frequent breeding appeared to be the time required to replace the wing and tail feathers.
Details are given of the breeding biology, all aspects of which were strongly influenced by frequent and unpredictable food shortages. The average incubation period was 49 days but this was prolonged by temporary desertion due to food shortages. Chick growth and fledging periods (62 to 100 days) were variable. There was no well–defined desertion period and birds were experimentally induced to feed young for up to 120 days.
Overall nesting success was low (26%) and adult survival between breeding attempts was high (92–95%).
Food, planktonic fish larvae and Crustacea, appeared to be rarely abundant and details are given of the effect of food shortage on the breeding.
The breeding synchrony was brought about by food shortages. The ultimate factor controlling breeding appeared to be the availability of food for egg formation and there was no possibility of birds timing breeding so that young were being fed at a time of food abundance.
A comparison is made of the breeding of eight Puffinus species.     

THE IMPORTANCE OF CERTAIN ASSEMBLAGES OF BIRDS AS "INFORMATION-CENTRES" FOR FOOD-FINDING

Evidence is presented to support the hypothesis that communal roosts, breeding colonies and certain other bird assemblages have been evolved primarily for the efficient exploitation of unevenly-distributed food sources by serving as "information-centres".
Predation-pressure is regarded as being the most important factor "shaping" the assemblages. The shaping involves the choice of inaccessible or otherwise safe sites, optimum dispersal, mutual awareness of attack and joint defensive tactics, and serves to minimise the vulnerability to predation which would otherwise result when birds mass together in conspicuous, and often predictable centres.     

Alternate Routes to Sociality in Jays--With a Theory for the Evolution of Altruism and Communal Breeding

The social systems of New World species of jays are comparedbehaviorally and ecologically. Two lines leading to socialityare identified, one connecting the pair-in-territory systemwith the colonial system and a second connecting the pair-in-territorysystem with the communal system. Ecological factors in the evolution of communal breeding andits associated altruistic behavior are considered mainly inthe genus Aphelocoma, but other birds are treated briefly. Threeprincipal origins of communality and altruism in birds are identified:from colonies, from a surplus of males, and by retention ofyoung in the family, the latter being of greatest importance.The theory proposed for the evolution of communal breeding andaltruism emphasizes three main processes: 1) K-selection, (2)kin selection, and (3) kin-group selection in the same chronologicalorder.     

THE PRIMARY MOULT OF WADERS ON THE ATLANTIC COAST OF MOROCCO

Data from 3659 waders of 23 species live-trapped in the years 1971-73 on the Atlantic coast of Morocco during the period of autumn moult and migration are analysed to estimate duration and timing of primary moult. Common Sandpiper was the only species to moult primaries in its first autumn (unless published ageing criteria are incorrect). Several species showed a low incidence of arrested primary moult and a higher incidence was observed in Ringed, Kentish and Grey Plovers. This is discussed in relation to breeding and migration. Similar rates of primary feather replacement relative to specific moult duration were observed in all species for which information was available. Comparisons between species and with published studies showed that variations in rate of moulting between species and between different geographical populations of the same species were largely due to differences in feather growth rate rather than in the numbers of primaries concurrently in growth. Variations in rate between individuals of the same population were achieved, at least in the first part of moult, by differences in feather dropping rate resulting in differences in the numbers of primaries growing concurrently. The timing and duration of moult in different populations and differences between breeding and non-breeding components were closely related to the requirements of other annual cycle activities, notably breeding and migration. Non-breeding birds summering in Morocco had started moult early. Locally breeding birds had an early start to a fairly slow moult which overlapped with breeding and which in some cases passed through an arrested stage. Birds breeding in cold temperate and arctic regions and wintering in Morocco moulted in a short time soon after arrival. In some cases, notably in Ringed Plovers, birds had commenced moulting on the breeding grounds and arrested moult during migration. Most Redshank and possibly Dunlin migrated in active wing moult. The fastest primary moult was achieved by high arctic breeding birds, Curlew Sandpiper and possibly Little Stint, which stopped to moult in Morocco before moving on to wintering areas further south. This situation is contrasted with that of populations of these two and other species wintering in the southern hemisphere where moult occurs over an extended period during the northern winter.     

Common Myna Roosts Are Not Recruitment Centres

We studied communal roosting in the Common Myna (Acridotheres tristis) in the light of the recruitment centre hypothesis and predation at the roost. The number and sizes of flocks departing from and arriving at focal roosts were recorded over a two year period. We also recorded the sizes and behaviour of foraging flocks. We found that flock sizes of birds departing from roosts at sunrise were larger than those at the feeding site, suggesting that there was no recruitment from the roosts. Flocks entering the roosts during sunset were larger on average than those leaving the following sunrise, suggesting no consolidation of flocks in the morning. Flocks entering the roosts at sunset were also larger on average than those that had left that sunrise, although there was no recruitment at the feeding site. There was no effect of group size on the proportion of time spent feeding. Contrary to expectation, single birds showed lower apparent vigilance than birds that foraged in pairs or groups, possibly due to scrounging tactics being used in the presence of feeding companions. Thus, the recruitment centre hypothesis did not hold in our study population of mynas. Predation at dawn and dusk were also not important to communal roosting: predators near the roosts did not result in larger flocks, and resulted in larger durations of arrival/departure contrary to expectation. Since flock sizes were smallest at the feeding site and larger in the evening than in the morning, but did not coincide with predator activity, information transfer unrelated to food (such as breeding opportunities) may possibly give rise to the evening aggregations.     

EXPERIMENTS ON THE LIMITATION OF BIRD NUMBERS BY TERRITORIAL BEHAVIOUR

(1) This paper examines removal and other experimental studies on the role of territorial behaviour in the limitation of bird densities. Experimental design is discussed, as are the types of conclusions that can be drawn. (2) Experiments have been conducted on more than 40 species from a wide range of taxonomic groups. Most provided evidence for density limitation, probably mediated by territorial behaviour, and for the existence of a non-territorial sector able to take territories when territory owners were removed or when additional habitat was made available. Experiments in the breeding season indicated that some surplus birds, although younger on average then territorial birds, were sexually mature and able to breed when given the chance. In some species, replacement birds bred less well than other territorial birds, but in other species no difference was apparent between the two groups. (3) Other experiments indicated that, while density was limited by territorial behaviour in good habitats, this was not the case in poor habitats, and that some individuals would move from poor to good habitat when territory owners were removed from good habitat. (4) In some species, non-territorial birds of breeding age were present in the population despite the existence of vacant territorial sites, while in others replacements were observed on good territories but not on poor ones. The implications were that site quality influenced whether settlement, defence and breeding occurred, and that some individuals had more stringent site requirements than others. In some seabirds' sites in the centre of a colony were more attractive than sites on the edges. (5) Many land-bird species that have been studied show two peaks of territorial activity each year, in autumn and in spring, and a limitation on breeding density can occur at either or both seasons, depending on conditions. (6) The fact that densities of many bird species fluctuate greatly from year to year is not inconsistent with the limitation of density by territorial behaviour. Several mechanisms are apparent through which density might be limited by territorialism at different levels in different years, so that surplus non-territorial birds are present in years of low territorial density, as well as in years of high territorial density. (7) Experiments have shown that density is limited by the presence of territorial birds, in some species at different levels in different areas or years. The next step is to find whether density is regulated by the presence of territorial birds over a period of years in a density-dependent manner. For this, observational data are required to find whether the proportion of birds that is excluded by territorial behaviour each year varies with the total number available for settlement. Such studies can be made only on species in which non-territorial birds can be counted accurately, as well as territorial ones.     

Survival in a long-lived territorial migrant: effects of life-history traits and ecological conditions in wintering and breeding areas

Despite its key role in population dynamics and evolutionary ecology, little is known about factors shaping survival in long‐lived territorial species. Here, we assessed several hypotheses that might explain variability in survival in a migratory Spanish population of a long‐lived territorial species, the Egyptian vulture Neophron percnopterus, using a 16‐year monitoring period and live‐encounter histories of 835 individually marked birds. Cormack‐Jolly‐Seber capture–recapture models showed no evidence for effects of sex or nestling body condition on survival. However, the normalized difference vegetation index (NDVI; an indicator of primary productivity) of natal territories had positive effects on juvenile survival, indicating that environmental conditions experienced early in life can determine survival prospects. Survival increased with age (0.73±0.02 in the first 2 years to 0.78±0.03 in years 3 and 4) to later decrease when birds were five years old (0.60±0.05), the age at which they acquire the adult plumage, abandon the communal lifestyle of juveniles, and may look for a breeding territory. At older ages, survival was higher for non‐breeding (0.75±0.02) and breeding adults (0.83±0.02). Among the latter, birds that recruited into better territories had higher survival prospects. Age‐specific variation in survival in this species may be related to behavioural changes linked to dispersal and recruitment into the breeding population, while survival prospects of adult birds strongly depend on breeding territory selection. These results suggest a tradeoff between recruiting soon, and thus reducing mortality costs of a long and extensive dispersal period, and trying to recruit into a good quality territory. Finally, annual survival rates for birds of all age classes were positively related with the NDVI in their African wintering grounds. Although this relationship was probably mediated by food availability, further research is needed to properly identify the limiting factors that are affecting trans‐Saharan migrants, especially in light of global climate change.     

THE SOCIABLE WEAVER,PART 3: BREEDING BIOLOGY AND MOULT

Maclean, G. L. 1973. The Sociable Weaver, Part 3: Breeding biology and moult. Ostrich 44: 219–240.

Rain or some associated phenomenon is the principal Zeitgeber releasing breeding in the Sociable Weaver. The species does not breed in the absence of rain. The same nest chambers are used for breeding as are used for roosting throughout the year. The Sociable Weaver is monogamous. The clutch size varies from two to six eggs, larger clutches being more common after good rains than in relatively poorer breeding periods. Food supply may therefore be the proximate factor regulating clutch size. Replacement clutches are not necessarily smaller than first clutches. The mean clutch size within a breeding period decreases with an apparent decrease in food supply. The parents share parental duties about equally. Up to four successive broods may be raised in a single breeding period; a breeding period may last up to nine months and may occur at any time of the year according to the somewhat erratic rainfall which averages about 226 mm per year in the study area.

First broods help their parents to feed later broods; fourth brood chicks may therefore be fed by as many as 11 birds (nine young and two parents). This has survival value especially toward the end of a breeding period when food is scarce. Of similar value is the habit of starting incubation with the first or second egg of the clutch; in a relatively poor season older chicks will survive while younger ones will starve, thereby effectively and quickly reducing brood size. Young birds moult into adult plumage at four months, but do not normally leave the home colony. The sexes are indistinguishable at all ages, but there is an approximate ratio of eight males to five females in the study area.

Wing moult is slow: each remex takes about a month for replacement. Body moult occurs within the space of a month, usually after rain while the birds are breeding. Primary remiges are moulted proximo-distally from 1 to 9; secondaries are moulted disto-proximally from 1 to 6. Body moult is antero-posterior with the dorsal surface slightly in advance of the ventral surface.     

BREEDING OF THE WHITE-BACKED AND RÜPPELL'S GRIFFON VULTURES, GYPS AFRICANUS AND G. RUEPPELLII

The breeding season of two species of griffon vultures are described. Rüppell's Griffon Vulture lays 2–3 months earlier than the White-backed Griffon. Young birds were hand-reared to determine their food requirements during growth; these estimates were combined with the food requirements of adult birds to make an estimate of the amount of food a parent bird needs to obtain when it is rearing young. The amount of food actually obtained by a group of birds was recorded from the size of the crops of birds returning to the breeding colony in the afternoon. The comparison of the estimates of the food obtained and the food required through the breeding season suggested that there may be a period during rearing when there was insufficient food available to satisfy the food requirements of both chick and adult. Chicks were found to have a very high survival rate and were probably receiving sufficient food. Presumably adult birds were not therefore receiving sufficient food, and the examination of a sample of adult birds for body condition through the breeding season showed a clear decline in their fat deposits. It was considered that in both species, breeding was timed so that the young left the nest at a period in the year when food conditions were good and the young birds could feed with little competition from adults. The parent birds therefore had to rear young during a season in the year when food conditions were not always adequate and they had to rely on utilising fat reserves. The food conditions for vultures during this study were probably favourable and during years of food shortage breeding may become impossible, or restricted to the most aggressive and dominant individuals.     

THE EFFECT OF AGE ON THE BREEDING BIOLOGY OF THE KITTTWAKE RISSA TRIDACTYLA

1. From 1954 to 1956 inclusive, the biology of individual marked Kittiwakes was studied at North Shields, Northumberland.
2. It was concluded that older Kittiwakes reacted to the breeding stimulus earlier, more intensively and with greater success than younger breeding birds.
3. Birds with previous breeding experience returned to the colony before birds breeding for the first time and these before non-breeders.
4. Before breeding started, birds which had bred previously spent more of their time at the colony than those about to breed for the first time.
5. Birds breeding for at least the second time laid the first egg 7·5 days earlier than those breeding for the first time.
6. Breeding started one day later for every four days the return to the colony was postponed.
7. Older breeding birds showed greater nest-site tenacity, laid larger clutches and had greater breeding success than younger birds.
8. The chicks in broods of two (but not of one) increased in weight more rapidly where the parents had previous experience.
9. Breeding Kittiwakes showed strong colony tenacity, but 24% of the marked non-breeding birds were subsequently seen in other colonies.
10. Over half the birds retained the same mate as in the previous year.