Plasticity of moult and breeding schedules in migratory European Stonechats Saxicola rubicola

Timing is crucial in seasonal environments. Passerine birds typically use a combination of physiological mechanisms and environmental cues to ensure that breeding, moult and migration occur without major temporal overlap and under the most favourable conditions. However, late in the breeding season some individuals initiate additional clutches , whereas others initiate moult. Such alternative strategies are thought to reflect trade‐offs between reproductive benefits and timely investment in maintenance and survival. The degree of seasonal plasticity differs between species, depending on the mechanisms that govern their annual routine. Migrants are generally under pressure to complete breeding and moult before the autumn departure and often show little plasticity. We studied seasonal plasticity of breeding and moult schedules in the European Stonechat Saxicola rubicola. This species, an obligate short‐distance migrant in Central Europe, sometimes initiates late clutches after typically at least two earlier breeding attempts. Based on life‐history theory and on observations in captivity, which revealed photoperiodic regulation of breeding and moult, we predicted relatively little seasonal plasticity in Stonechats. We further predicted that reproductive gains of late breeders should be offset by reduced survival. These predictions were tested on long‐term field data, using Underhill–Zucchini models to estimate moult. Late breeding occurred in c. 40% of pairs and increased their reproductive success by a third. Both sexes modified moult timing but in different ways. Late breeding females postponed moult approximately until chick independence without compensating for delay by faster moult. Males started moult on time and overlapped it with breeding, associated with markedly slowed plumage change. Sex differences in moult score increased with lay date, but due to their respective modifications, both sexes delayed moult completion. Nonetheless, we could not detect any evidence for survival costs of late breeding. Breeding and moult of European Stonechats appear relatively flexible, despite migratory schedules and photoperiodic programs for seasonal timing. Individuals can modify seasonal behaviour in late summer, presumably depending on their condition, and may profit considerably from extended breeding.     

Weak breeding seasonality of a songbird in a seasonally arid tropical environment arises from individual flexibility and strongly seasonal moult

In some tropical birds, breeding seasonality is weak at the population level, even where there are predictable seasonal peaks in environmental conditions. It therefore remains unclear whether individuals are adapted to breeding at specific times of the year or flexible to variable environmental conditions. We tested whether the relative year‐round breeding activity of the Common Bulbul Pycnonotus barbatus arises due to within‐individual variability in breeding dates. We collected data from 827 birds via mist‐netting over 2 years with corresponding local weather data. We used a combination of climate envelope and generalized linear mixed models to explore how the timing of breeding is influenced by time of year, individual variation, rainfall and temperature in a West African savannah where seasonal precipitation determines annual variation in environmental conditions. We also pooled 65 breeding records from 19 individuals recorded between 2006 and 2017 based on brood patch occurrence and behavioural observation to compare within‐individual and population variability in breeding dates. We show that the breeding dates of individuals may be as variable as for the population as a whole. However, we observed a seasonal peak in juvenile occurrence that varies significantly between years. Models suggest no relationship between nesting and moult, and within‐year variation in rainfall and temperature, and birds were unlikely to breed during moult but may do so afterwards. Moult was very seasonal, correlating strongly with day length. We suggest that because environmental conditions permit year‐round breeding, and because reproductive output is subject to high predation risk, there is probably a weak selection for individuals to match breeding with variable peak conditions in the environment. Instead, moult, which always occurs annually and successfully, is probably under strong selection to match variable peak conditions in the environment so that long‐term survival ensures future reproduction.     

Seasonal breeding, moulting and weight changes among birds of dry deciduous forest in North India

The timing of breeding among Indian birds varies between species and also regionally within species. Marked differences can occur between adjacent areas, depending on the seasonal availability of moisture for plant growth. Birds of dry deciduous woodland have two peaks of breeding activity, in spring (March-April) and during the rains (July-September), with moult following the second peak in most cases. Rates of moult vary greatly between species, but practically all terminate moult by the end of November and this may truncate the potential breeding season for some species.
Changes in weight show a marked contrast with seasonal patterns observed in temperate latitudes, weights being inversely correlated to mean temperature. It is suggested that, in the case of some species which occupy a wide range of vegetation types, observed patterns of breeding and moulting are a compromise between the optimum strategies for different habitats.     

Annual cycle of the Helmeted Honeyeater Lichenostomus melanops cassidix, a sedentary inhabitant of a predictable environment

The annual cycle of breeding, moult and weight variation in the Helmeted Honeyeater Lichenostomus melanops cassidix , a sedentary bird of temperate southeast Australia, is documented. Breeding and moult were sequential unimodal annual events, whose timing was highly consistent between years. However, overlap of breeding and moult was frequent, and some individuals even commenced primary moult before laying their final clutch. The timing of the post-juvenile moult was coincident with that of adults. Early-hatched young moulted within a few months of hatching, but late-hatched young deferred moult for a year. Helmeted Honeyeaters were heaviest in autumn and early winter, and lightest in spring and early summer, a cycle most consistent with the redirection of all available resources to reproduction. The long breeding season (seven-and-a-half months) of the Helmeted Honeyeater, extensive overlap of breeding and moult, and other life-history attributes including small clutch size, are more consistent with the described bio-rhythmic patterns for birds in the humid tropics than the temperate zone. However, the Helmeted Honeyeater has a fairly rapid primary moult rate, unusual amongst species that overlap moult and breeding. This combination of attributes reflects the stable, somewhat seasonal environment occupied and the resource monopoly established by this tightly territorial subspecies. We speculate that extension of the breeding season, by overlapping breeding and moult, is one of the few options available to vary life-history strategies amongst 'old-endemic' Australian birds of the temperate zone.     

Optimal moult strategies in migratory birds

Avian migration, which involves billions of birds flying vast distances, is known to influence all aspects of avian life. Here we investigate how birds fit moult into an annual cycle determined by the need to migrate. Large variation exists in moulting patterns in relation to migration: for instance, moult can occur after breeding in the summer or after arrival in the wintering quarters. Here we use an optimal annual routine model to investigate why this variation exists. The modelled bird's decisions depend on the time of year, its energy reserves, breeding status, experience, flight feather quality and location. Our results suggest that the temporal and spatial variations in food are an important influence on a migratory bird's annual cycle. Summer moult occurs when food has a high peak on the breeding site in the summer, but it is less seasonal elsewhere. Winter moult occurs if there is a short period of high food availability in summer and a strong winter peak at different locations (i.e. the food is very seasonal but in opposite phase on these areas). This finding might explain why only long-distance migrants have a winter moult.     

Moult of Malayan Barn Owls Tyto alba

Moult in Malayan Barn Owls Tyto alba was studied in two pairs of wild collected captive birds and from feathers taken from nest sites throughout peninsular Malaysia.
Post-juvenile captive birds moulted nearly to completion prior to first breeding, beginning with P6 at a mean age of 301.5 days. This contrasted with the only other study of moult in captive Barn Ow-Is in Germany when moult began at an age of 400 days, and then continued for a protracted period of two years separated by a suspension of moult during the normal breeding season.
The complex sequence of moult in primaries and secondaries both in the Malayan and German birds was similar.
Moult among adult Malayan birds in the wild showed a broad and somebyhat irregular seasonal trend With lower incidence during peak breeding periods.     

Seasonal colour change by moult or by the abrasion of feather tips: a comparative study

Many birds undergo seasonal changes in plumage coloration by prebreeding moult, abrasion of cryptic feather tips, or both. Seasonal dichromatism is thought to result from optimizing coloration to the conflicting demands of different life-cycle periods, sexual selection for conspicuousness being substantial during the mating season, whereas selection for camouflage and for social signals may act in all seasons. Furthermore, energetic and time demands may constrain the extent of moult, thereby limiting colour change. We investigated the relative importance of several factors in shaping this variation in a songbird clade using phylogenetic comparative methods. We found that prebreeding moult relates most strongly to breeding onset and winter diet, demonstrating that both time and food availability constrain feather replacement. Feather abrasion was best predicted by winter flocking behaviour, and secondarily by open habitats, implying that exposure to predators and the simultaneous need for social signalling may favour the expression of partially obscured ornaments in the non-breeding season. The combined occurrence of prebreeding moult and feather abrasion was associated with the polygynous mating system, suggesting that species under strong sexual selection may employ both strategies of colour change to ensure the full expression of breeding coloration.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 711–721.     

Annual routines of non-migratory birds: optimal moult strategies

In a periodically changing environment it is important for animals to properly time the major events of their life in order to maximise their lifetime fitness. For a non-migratory bird the timing of breeding and moult are thought to be the most crucial. We develop a state-dependent optimal annual routine model that incorporates explicit density dependence in the food supply. In the model the birds' decisions depend on the time of year, their energy reserves, breeding status, experience, and the quality of two types of feathers (outer and inner primaries). Our model predicts that, under a seasonal environment, feathers with large effects on flight ability, higher abrasion rate and lower energetic cost of moult should be moulted closer to the winter (i.e. later) than those with the opposite attributes. Therefore, we argue that the sequence of moult may be an adaptive response to the problem of optimal timing of moult of differing feathers within the same feather tract. The model also predicts that environmental seasonality greatly affects optimal annual routines. Under high seasonality birds breed first then immediately moult, whereas under low seasonality an alternation occurs between breeding and moulting some of the feathers in one year and having a complete moult but no breeding in the other year. Increasing food abundance has a similar effect.     

THE RELATIONSHIP BETWEEN CLIMATE AND ANNUAL CYCLES IN THE COTINGIDAE

This survey is based primarily or the state of moult of over 4000 specimens of cotingas from all parts of the neotropical region. The seasonality of moult thus revealed is combined with existing knowledge of breeding seasons and seasonal environmental changes in an attempt to work out the broad pattern of annual cycles and their relation to climate. Within any local population the date of onset of moult may vary according to sex and age. In genera in which both sexes participate in nesting, males and females begin to moult at about the same time, or the males slightly in advance of the females. In genera with marked sexual dimorphism, in which only the female attends the nest, males may begin to moult well before females, at about the time that the latter begin egg-laying. The former group includes the genera Pachyramphus and Tityra, comprising species that are largely insectivorous, and the latter group includes the more specialized frugivorous genera. In all areas with well-marked seasonality, the ‘frugivorous group’ moults on average before the ‘Pachyramphus group’. It appears to be a general rule for first-year birds to moult earlier than older birds. A regional survey embracing all parts of the neotropical region shows that the peak of onset of moult occurs towards the end of the dry season (frugivorous group) or early in the wet season (Pachyramphus group). The changing moult seasons, strikingly in conformity with the geographical changes in the period of heaviest rainfall, are traced along a number of transects from Mexico in the north to Paraguay and Bolivia in the south. Such evidence as there is suggests that the main period of onset of moult in the frugivorous and Pachyramphus groups coincides with the period when their food is approaching or at its seasonal peak of abundance. It seems that both breeding and moult, which are almost entirely mutually exclusive, are as far as possible timed to coincide with this most favourable period; but whereas the moult takes a more or less fixed length of time the period when breeding is possible varies greatly in different species. Widely different patterns of annual cycle may result from the interaction of the two processes. Examples are given both from the cotingas and from species of other families with similar ecology. The proximate factors controlling the timing of the moult are briefly considered. It is suggested that increasing food availability is the main environmental controlling factor, and that an endogenous circannual cycle of moult must also be involved.     

Seasonal stress physiology and body condition differ among co-occurring tropical finch species

Seasonal changes in avian hormonal stress responses and condition are well known for common species found at temperate and arctic latitudes, but declining and tropical species are poorly studied. This study compares stress and condition measures of co-occurring declining and non-declining tropical grass finch species in Australia. We monitored declining Gouldian finches (Erythrura gouldiae) and non-declining long-tailed and masked finches (Poepila acuticauda and P. personata) during two seasons that are potentially stressful: peak breeding (early dry season when food is plentiful) and moult (late dry to early wet season when food may be scarce). We measured body condition (muscle and fat), haematocrit, and stress response to capture using plasma corticosterone and binding globulin concentrations. All species had higher muscle and lower fat indices during breeding than moult. Haematocrit did not consistently differ between seasons. Long-tailed finches had higher stress responses during breeding than moult, similar to other passerines studied. Masked finches showed no seasonal changes in stress response. Gouldian finches had stress response patterns opposite to those of long-tailed finches, with higher stress responses during moult. However, seasonal trends in Gouldian and long-tailed finch stress responses sometimes differed between years or sites. The differences in stress response patterns between species suggest that the declining Gouldian finch is more sensitive to recent environmental changes which are thought to further reduce grass seed food resources during the late dry to early wet season. Retention of stress responsiveness during a protracted moult could increase the survival potential of Gouldian finches. This study highlights the utility of stress and condition indices to determine the sensitivity of co-occurring species to environmental conditions.     

A comparison of breeding and moult cycles and life histories in two tropical starling species: the Blue-eared Glossy Starling Lamprotornis chalybaeus and Rüppell's Long-tailed Glossy Starling L. purpuropterus

The Blue-eared Glossy Starling Lamprotornis chalybaeus and Rüppell's Long-tailed Glossy Starling Lamprotornis purpuropterus were investigated in the field and in aviaries at Lake Nakuru National Park, Kenya for seasonality in reproductive activity and moult. The former species was found to be a seasonal breeder which nests after the onset of the heavy rains in April. Although some birds had large gonads prior to the rains in the dry season no nesting occurred. The rains were contemporary with increases in gonadal size and the plasma titres of LH, testosterone (T) in males and estradiol (E2) in females. These hormones are associated with the initiation of breeding activity. As breeding ceased in July and the moult began, the plasma titres again decreased. There was a bimodal breeding pattern which paralleled a change in biotope preference for nesting. Early nests, in the heavy rains, were on the open savanna and later nests were in the acacia forest. Late nesting birds also had delayed peaks in gonadal size, plasma titres of LH, T and E2 and a delayed moult onset. Data on individual captive birds demonstrate that these annual cycles have a distinctly individual character superimposed on the seasonal trends. In Rüppell's Long-tailed Glossy Starlings no seasonality in breeding was found although the flight feather moult commenced and was completed in all individuals at about the same time. The moult extended over about ten months, so a great deal of breeding-moult overlap was present. The absence of seasonality in field birds was reflected in the aviary birds, which had no pronounced cycles in the reproductive parameters measured (gonadal size, LH, T and E2 plasma titres). Breeding in field birds was regulated on a pair basis and correlated with increases in duetting. The striking differences in the seasonal organization between this species and Blue-eared Glossy Starlings were presumably due to the different biotope preferences and social behaviour of the two species.     

A twelve-month field study of the West African thrush Turdus pelios (Passeriformes: Muscicapidae). Part 2: annual cycles

In Africa, birds inhabiting forested regions are less seasonal in their activities than those from open areas. In order to study annual cycles in forest regions of South western Nigeria, West African Thrushes (Turdus pelios) were mist-netted and banded during the last two weeks of each month. The nest is a cup-shaped structure built out of grasses, herbs, weeds, roots and earth laid out in a clockwise manner. Only the nesting tree and feeding sites were defended during the breeding period. The clutch size was 2.69 +/- 0.20 eggs with a mean incubation period of 14.11 +/- 0.26 days. The mean nestling period was 15 +/- 1.00 days. The nestlings were fed on a variety of plant and animal matter, of which grass seeds and insects were predominant. Moult was found to be protracted with a population moult period of 194 days and a much shorter individual moult period. Moult and breeding periods were spread out: moult period dovetailed into the breeding period. The birds were found to gain weight during the period but they attained their maximum weight in August after the moult period. The lowest weight was recorded in February, during the peak of the dry season, when food availability was lower.     

Ecological correlates in the evolution of moult strategies in Western Palearctic passerines

Moult is an important process in the life cycle of birds. Passerines differ widely in the number, seasonality and extension of moult episodes, but the incidence of birds ecology on this variation remains largely uninvestigated. We analysed the patterns of moult in European passerines in relation to their distribution, migration and sexual dichromatism. Longer migrations and southern wintering quarters were characteristic of species with complete moults in summer and an additional moult in winter. The main moult in species with larger seasonal changes in sexual dimorphism tended to be scheduled just before the start of the breeding season, suggesting a link between sexual selection and the timing of moult. These patterns strongly support the importance of migration and dichromatism on the evolution of moult strategies.     

Breeding seasons of sea-birds in the Galapagos Islands

The sea-birds breeding in the Galapagos Islands show a diversity of breeding cycles. Some species have rigidly fixed annual breeding while others breed throughout the year but have peaks of breeding at less than annual intervals. The eight species which have non-annual breeding are probably breeding as often as possible with the interval between the end of a breeding attempt and the start of the next being the time needed to moult the wing and tail feathers. Only one species is definitely known to breed and moult at the same time.
Although there are well marked seasonal fluctuations in the sea temperature, regular sampling failed to demonstrate any regular fluctuations in the surface plankton. The available evidence suggests that food for some sea-birds is erratic and unpredictable. Some non-annual breeding species have their breeding synchronized by severe food shortages which delay breeding, presumably because females cannot find enough food to form eggs, until conditions improve.
Timing of the breeding season in annual breeders is less easily explained but some species may be feeding well away from the islands in areas where there is a regular fluctuation in the food supply. Most of the annual breeders have prolonged breeding seasons and in two species breeding is out of phase on different islands. Perhaps species are influenced by some weak annual variation in food supply which makes it disadvantageous to breed in a few months of the year.     

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.     

THE BREEDING SEASON OF BIRDS IN THE LOWLAND RAINFOREST AND IN THE MONTANE FOREST OF WEST CAMEROON

An analysis is made of over a thousand precise bird breeding records for a forest area in West Cameroon. In the lowland forest there is no universal breeding season although most families have a preferred nesting season. On the other hand. in the montane forest breeding is seasonal and takes place in the drier months. These conclusions are supported by field observations and by the examination of over 5000 birds for gonad state, plumage wear and moult.
The remarkable difference in seasonality between the lowland and montane avifaunas is discussed with reference to climate, particularly to the mist which envelops the montane forest in the rainy season.     

Moult in Black-browed and Grey-headed Albatrosses Diomedea melanophris and D. chrysostoma

We recorded the age of individual wing and tail feathers of Black-browed and Grey-headed Albatrosses Diomedea melanophris and D. chrysostoma of known age and breeding status at Bird Island, South Georgia. Breeders and non-breeders of both species moult their rectrices annually. Non-breeders moult primaries biennially. In the first year of a cycle, the outer three and some inner primaries are moulted descendantly; in the next year the inner primaries are moulted ascendantly, starting from primary seven. There is a general progression to moulting equal numbers of primaries in each half of the cycle by the time breeding starts at about 10 years of age. Grey-headed Albatrosses usually moult fewer primaries than Black-browed Albatrosses, particularly as 3-year-olds, when they undertake substantial plumage change in body moult. Most secondaries in Black-browed Albatrosses have been replaced once by age 4 years. Breeding Black-browed Albatrosses continue the moult pattern established as immatures whether they fail or not, as do failed Grey-headed Albatrosses. Successful Grey-headed Albatrosses, which breed again 16 months later, moult their three innermost primaries after breeding in the remainder of the current year and, after a period when moult is interrupted, renew the remaining primaries the following year. Comparisons between species and between failed and successful birds within species indicate that moult rate is not closely linked to the length of the interval between breeding attempts. Interspecies differences are better explained by breeding latitude, with tropical albatrosses moulting twice as fast as sub-Antarctic species, possibly reflecting food availability outside the breeding season.     

Flexibility in the timing of post-nuptial moult among Red-billed Queleas Quelea quelea in Botswana in relation to the timing of breeding

The timing of primary moult of adult Red-billed Queleas Quelea quelea, captured as they were completing an unusually late breeding attempt at Francistown, northern Botswana, in June 2004, was compared with the timing of moult of birds breeding earlier in the season in north-west Botswana during two earlier years, 1971 and 1972. Differences between years in the dates when local colonies finished breeding (mid-March to late June) and between two localities in the same year (mid-March and late May) were matched by corresponding differences in the estimated dates of moult onset, ranging from mid-April to mid-June. Flexibility in the timing of moult among Red-billed Queleas in southern Africa evidently enables birds to take advantage of unusually late breeding opportunities by delaying moult onset and overlapping moult and breeding at the end of the nesting cycle. Such flexibility may also include moult interruption to permit late breeding, although its incidence in southern Africa is apparently low.     

The effects of long‐distance migration on the evolution of moult strategies in Western‐Palearctic passerines

Although feathers are the unifying characteristic of all birds, our understanding of the causes, mechanisms, patterns and consequences of the feather moult process lags behind that of other major avian life‐history phenomena such as reproduction and long‐distance migration. Migration, which evolved in many species of the temperate and arctic zones, requires high energy expenditure to endure long‐distance journeys. About a third of Western‐Palearctic passerines perform long‐distance migrations of thousands of kilometres each year using various morphological, physiological, biomechanical, behavioural and life‐history adaptations. The need to include the largely non‐overlapping breeding, long‐distance migration and feather moult processes within the annual cycle imposes a substantial constraint on the time over which the moult process can take place. Here, we review four feather‐moult‐related adaptations which, likely due to time constraints, evolved among long‐distance Western‐Palearctic migrants: (i) increased moult speed; (ii) increased overlap between moult and breeding or migration; (iii) decreased extent of plumage moult; and (iv) moult of part or all of the plumage during the over‐wintering period in the tropics rather than in the breeding areas. We suggest that long‐distance migration shaped the evolution of moult strategies and increased the diversity of these strategies among migratory passerines. In contrast to this variation, all resident passerines in the Western Palearctic moult immediately after breeding by renewing the entire plumage of adults and in some species also juveniles, while in other species juvenile moult is partial. We identify important gaps in our current understanding of the moult process that should be addressed in the future. Notably, previous studies suggested that the ancestral moult strategy is a post‐breeding summer moult in the Western Palearctic breeding areas and that moult during the winter evolved due to the scheduling of long‐distance migration immediately after breeding. We offer an alternative hypothesis based on the notion of southern ancestry, proposing that the ancestral moult strategy was a complete moult during the ‘northern winter’ in the Afro‐tropical region in these species, for both adults and juveniles. An important aspect of the observed variation in moult strategies relates to their control mechanisms and we suggest that there is insufficient knowledge regarding the physiological mechanisms that are involved, and whether they are genetically fixed or shaped by environmental factors. Finally, research effort is needed on how global climate changes may influence avian annual routines by altering the scheduling of major processes such as long‐distance migration and feather moult.     

Annual cycles in the African stonechat Saxicola torquata axillaris and their relationship to environmental factors

Seasonal changes in gonadal size, body weight, male song and the plasma titres of luteinizing hormone (LH), oestradiol and testosterone were investigated in a free-living population of African stonechats, Suxicola torquata axillaris , on Menengai Crater, Nakuru, Kenya. These were compared with the occurrence of nesting and moult in the population and a number of seasonally changing environmental factors including photoperiod, insect abundance and rain. Annual cycles in these stonechats appears to be regulated by endogenous events which are responsive to environmental factors on a seasonal basis. The dry season may play an important rôle in the synchronization of the population by inhibiting nesting in birds otherwise capable of reproductive activity. Rain is postulated as a time-giving cue for moult and gonadal regression. The possibility of photoperiodic changes at the equator acting as seasonal time-giving cues remains open.