Trade-off between reproduction and moult – a comparison of three Fennoscandian pied flycatcher populations

Organisms that reproduce at high latitudes are assumed to have evolved several adaptations to the short summer. For birds, and especially for long-distance migrants, there is a time constraint because both reproduction and moult must be completed before autumn migration. It has therefore been assumed that birds at northern latitudes must initiate their moult during reproduction more often than birds at low latitudes. To investigate how passerine birds breeding at different latitudes allocate their time between reproduction and moult, we compared timing of these activities during three consecutive breeding seasons in three widely separated populations of the pied flycatcher Ficedula hypoleuca. Our results show that the frequency of individuals with moult-breeding overlap, and moult initiation in relation to breeding stage, varied considerably among populations and years. In all three populations, female moult initiation was restricted to the late nestling period. The males had a more pronounced moult-breeding overlap than the females, but its duration was similar in all three study areas. Thus, there was no evidence for a more pronounced moult-breeding overlap at high compared with low latitudes. These results suggest that pied flycatchers sometimes accept a moult-breeding overlap, but that the time gained by having too extensive an overlap between reproduction and moult does not outweigh the associated costs. Long-distance migrants breeding at northern latitudes apparently experience a trade-off between reproduction and somatic investment during moult. We therefore suggest that a pronounced moult-breeding overlap is not a typical strategy used by long-distance migrants to adjust to the short breeding season at northern latitudes. Received: 7 May 1998 / Accepted: 24 August 1998     

Do captive waterfowl alter their behaviour patterns during their flightless period of moult?

Many different behavioural changes have been observed in wild waterfowl during the flightless stage of wing moult with birds frequently becoming inactive and reducing time spent foraging. Increased predation risk, elevated energetic demands of feather re-growth and restriction of foraging opportunities are thought to underlie these changes. By studying captive populations of both a dabbling and a diving duck species at the same site, we determined whether captive birds would reflect the behavioural responses of wild waterfowl to moult. The time-budgets of 42 Common Eiders, Somateria mollissima, (a diving duck) and 18 Garganeys, Anas querquedula, (a dabbling duck) were recorded during wing moult (July–August) and non-moult (January) with behaviour recorded under six categories. Despite captivity providing a low predation risk and constant access to food, birds altered their behaviour during the flightless period of wing moult. Time allocated to foraging and locomotion decreased significantly during moult compared to non-moult periods, while resting time increased significantly. Moulting Eiders underwent a greater reduction in time spent foraging and in locomotion compared with Garganeys, which is likely to be in response to a higher energetic cost of foraging in Eiders. It is possible that increased resting in both diving and dabbling ducks reduces their likelihood of detection by predators, while allowing them to remain vigilant. We demonstrate that there is much potential for using captive animals in studies that can augment our knowledge of behaviours of free-living conspecifics, the former being a hitherto under-exploited resource.     

The ‘chaotic’ flight feather moult of the Steppe Buzzard Buteo buteo vulpinus

Steppe Buzzards breed in Eurasia and spend the non-breeding season in Africa. Adults moult some flight feathers during the breeding season and some during the non-breeding season. Moult is arrested during migration. The extent of moult of flight feathers in adults is highly variable between individuals in southern Africa, with the renewal of two primaries, three secondaries and five rectrices as the most frequently encountered pattern. Time spent on the non-breeding grounds in South Africa is too short to allow for a sequential moult. Moult of flight feathers is restricted to the almost synchronous dropping of a number of feathers upon arrival, with few being replaced subsequently. Any of the flight feathers can be replaced in southern Africa, and the pattern of renewal in primaries and secondaries cannot be distinguished from random. Tail feathers are replaced in an alternating (transilient) pattern. Moult in the non-breeding areas may primarily be complementary to moult on the breeding grounds, but these two partial moults per year are insufficient to renew all flight feathers annually. Middle secondaries and central tail feathers are regularly carried over to a third moult, but this is rare for primaries.     

Rapid moult among palaearctic passerines in West Africa- an adaptation to the oncoming dry season?

Moult of some palaearctic migrant passerines was studied at a wintering site in northern Ghana, West Africa. We present data on moult pattern and estimated moult duration for a sample of species. Savi's Warbler Locustella luscinioides , Woodchat Shrike Lanius senator and probably Grasshopper Warbler Locustella naevia deviated from the ordinary descendent moult sequence. Different measures of moult speed (moult speed of retrapped individuals, growth of individual feathers, wing raggedness and first date of completion of moult) showed unexpectedly rapid moult among individuals of several species, notably the Great Reed Warbler Acrocephalus arundinaceus and the Sedge Warbler A. schoenobaenus. The rapid moult might be an adaptation to a limited period of abundant food which comes to an end at the beginning of the dry season.     

Primary moult of continental Black-tailed Godwits Limosa limosa limosa in the Doñana wetlands,Spain

Capsule: Most Continental Black-tailed Godwits Limosa limosa limosa using the Doñana wetlands during post-breeding migration appear to begin moult before they arrive and suspend moult before they migrate onwards to West Africa.

Aims: We aim to describe the primary moult strategies and patterns in the Continental Black-tailed Godwits using the Doñana wetlands, a major passage and wintering area for waterbirds in southern Spain.

Methods: Individual godwits were captured, marked and their primary moult was scored in Doñana during the non-breeding season (June–March) in 2011 and 2012. Data from resightings of colour-marked godwits and birds equipped with satellite transmitters were used to estimate stopover duration during post-breeding migration (June–September) to determine if godwits move to West Africa before completing their primary moult.

Results: Average primary moult duration was estimated to be 84 days?±?9 se, during 29 June–21 September and did not differ between sexes. Only 2% of individuals were observed with suspended moult. We estimated stopover duration in Doñana to be 13 days?±?2 se before migrating to West Africa.

Conclusions: Most godwits stage for about two weeks in the Doñana wetlands during southward migration, moult their primaries and appear to suspend moult before crossing the Sahara. Others may complete their primary moult in Doñana, or elsewhere in Europe and overwinter in Doñana where increasing numbers of godwits have been detected in recent years. A few individuals may finish the moult in Doñana and migrate to West Africa late in the post-breeding season.     

Why renew fresh feathers? Advantages and conditions for the evolution of complete post‐juvenile moult

Juveniles of several passerine species renew all of their fresh juvenile feathers immediately after fledging (complete post‐juvenile moult), in contrast to the majority, which perform a partial post‐juvenile moult. To understand the adaptive roles of this phenomenon we compared the quality of juvenile plumage in species that perform a complete post‐juvenile moult with that of species which perform a partial post‐juvenile moult; we similarly compared juveniles and adults in each of these groups. The quality of feathers was measured by mass of primaries, colour, and length. In species which perform a complete post‐juvenile moult the plumage quality of second‐year individuals, in their first breeding season, is similar to the plumage quality of adults, unlike those species that perform a partial post‐juvenile moult. In species which perform complete post‐juvenile moult, the quality of the feathers grown in the nest is lower than the quality of adult post‐breeding feathers. In contrast, in species which perform partial post‐juvenile moult the quality of the feathers grown in the nest is similar to that of adult post‐breeding feathers. We found that a complete post‐juvenile moult strategy is much more common 1) in residents and short‐distance migrants than in long‐distance migrants, 2) in southern latitudes, 3) in species with medium body mass and 4) in omnivores and granivores. Our results indicate two adaptive roles of the complete post‐juvenile moult strategy: 1) achieving high quality plumage in the first year which may increase individual survival probability and fitness and 2) allocating fewer resources to nestling plumage and more to nestling development, which enables the nestlings to leave the nest earlier, thus reducing the probability of encountering nest predators. We suggest that the complete post‐juvenile moult, immediately after fledging, is an optimal strategy in favourable habitats and under low time constraints, as in some tropical ecosystems.     

Phenotypic flexibility of a southern African duck Alopochen aegyptiaca during moult: do northern hemisphere paradigms apply?

Phenotypic flexibility during moult has never been explored in austral nomadic ducks. We investigated whether the body condition, organ (pectoral muscle, gizzard, liver and heart) mass and flight‐feather growth Egyptian geese Alopochen aegyptiaca in southern Africa show phenotypic flexibility over their 53‐day period of flightless moult. Changes in body mass and condition were examined in Egyptian geese caught at Barberspan and Strandfontein in South Africa. Mean daily change in primary feather length was calculated for moulting geese and birds were dissected for pectoral muscle and internal organ assessment. Mean body mass and condition varied significantly during moult. Body mass and condition started to decrease soon after flight feathers were dropped and continued to do so until the new feathers were at least two‐thirds grown, after which birds started to regain body mass and condition. Non‐moulting geese had large pectoral muscles, accounting for at least 26% of total body mass. Once moult started, pectoral muscle mass decreased and continued to do so until the flight feathers were at least one‐third grown, after which pectoral muscle mass started to increase. The regeneration of pectoral muscles during moult started before birds started to gain overall body mass. Gizzard mass started to increase soon after the onset of moult, reaching a maximum when the flight feathers were two‐thirds grown, after which gizzard mass again decreased. Liver mass increased significantly as moult progressed, but heart mass remained constant throughout moult. Flight feather growth was initially rapid, but slowed towards the completion of moult. Our results show that Egyptian geese exhibit a significant level of phenotypic flexibility when they moult. We interpret the phenotypic changes that we observed as an adaptive strategy to minimize the duration of the flightless period. Moulting Egyptian geese in South Africa undergo more substantial phenotypic changes than those reported for ducks in the northern hemisphere.     

Sexual dimorphism,moult and body condition of seabirds killed by longline vessels around the Prince Edward Islands, 1996–97

Ryan, P.G. 1999. Sexual dimorphism, moult and body condition of seabirds killed by longline vessels around the Prince Edward Islands, 1996–97. Ostrich 70 (3&4): 187–192.

A total of 393 seabirds from nine species killed by longline fishing in South African waters around the Prince Edward Islands during summer 1996–97 were sexed, measured, and their body fat levels and primary moults scored. Males of Whitechinned Petrels Procellaria aequinoctialis and Greyheaded and (Indian Ocean) Yellownosed Mollymawks Thalassarche chrysostoma and T. [chlororhynchosj bassi averaged 1–9% larger than females. Measures of bill depth were the most dimorphic characters in all three species. I recommend that if only a single measure is taken to estimate sex of adults, bill depth at the base be the standard measure for petrels, and bill depth at the nail be the standard measure for mollymawks. Discriminant functions are described for predicting the sex of unknown individuals; the proportion of misclassified birds ranged from 2–10%. With the exception of giant petrels Macronectes spp., only two young birds were actively moulting their primary flight feathers. Analysis of wear patterns among adult Greyheaded Mollymawk primaries conforms with studies of moult in this species at South Georgia. Fat scores did not differ between sexes. They tended to decrease during the breeding season, but trends were masked by great inter-individual variation. There was no significant relationship between fat scores and the presence of ingested plastics. Plastic debris was found in the stomachs of 90 birds from five species. Incidence in Whitechinned Petrel stomachs (37%) was less than that reported from birds collected off South Africa in the early 1980s (57%), but this is probably due to the predominance of breeding adults. The frequency and size of plastic loads decreased through the breeding season, which is consistent with the hypothesis that inter-generational transfer is important in the dynamics of ingested debris.     

The intermediate moult (First Ecdysis) of Schistocerca gregaria (Forskål) (Insecta,Orthoptera)

Ecdysis of the embryonic cuticle has been studied in detail. The initiation and mechanical processes involved in this ecdysis, and the associated differential size increases of cuticular parts have been given most attention. The roles of air swallowing and of thoracic contraction are most imoprtant for rupturing the old cuticle, while abdominal contractions assist the whole process. Some specialized cuticular structures are described.     

Mineral in skeletal elements of the terrestrial crustacean Porcellio scaber: SRμCT of function related distribution and changes during the moult cycle

Terrestrial isopods moult first the posterior and then the anterior half of the body, allowing for storage and recycling of CaCO₃. We used synchrotron-radiation microtomography to estimate mineral content within skeletal segments in sequential moulting stages of Porcellio scaber. The results suggest that all examined cuticular segments contribute to storage and recycling, however, to varying extents. The mineral within the hepatopancreas after moult suggests an uptake of mineral from the ingested exuviae. The total maximum loss of mineral was 46% for the anterior and 43% for the posterior cuticle. The time course of resorption of mineral and mineralisation of the new cuticle suggests storage and recycling of mineral in the posterior and anterior cuticle. The mineral in the anterior pereiopods decreases by 25% only. P. scaber has long legs and can run fast; therefore, a less mineralised and thus lightweight cuticle in pereiopods likely serves to lower energy consumption during escape behaviour. Differential demineralisation occurs in the head cuticle, in which the cornea of the complex eyes remains completely mineralised. The partes incisivae of the mandibles are mineralised before the old cuticle is demineralised and shed. Probably, this enables the animal to ingest the old exuviae after each half moult.     

Experiments on the physiology of southern and northern krill,Euphausia superba and Meganyctiphanes norvegica,with emphasis on moult and growth – a review

Physiological data are needed for life history studies on krill, and as parameters for input into energy budgets and models. In conjunction with moult and growth data, these may also prove useful for assessing the fishable biomass of krill. Here, the development of physiological concepts in experimental krill research is briefly evaluated, with emphasis on the gaps to be filled. Krill growth is very flexible, as well as strongly temperature and nutrition dependent. The polar Antarctic krill Euphausia superba grows as fast as the boreal species Meganyctiphanes norvegica, at least during the first 2.5 years, and the species are comparable in terms of physiological plasticity. Accordingly, as krill appear to adjust quickly to specific laboratory conditions, short-term experiments are essential if field conditions are to be reflected as closely as possible. Furthermore, direct comparisons between laboratory experiments and swarming studies in the field are advantageous. For these, M. norvegica is particularly well-suited, as swarms can be followed over longer times and more easily than in E. superba. For example, processes of moult and reproduction were found to be highly coordinated in swarms and populations of Northern krill. For this species a conceptual model of reproduction was developed based on a combination of short-term laboratory observations coupled with field data on moult and ovary stages. In further physiological experiments krill should be studied as groups when swarming. Using proxies, that is applying physiological and/or biochemical methods side by side, is a promising way to enhance the reliability of life history data.     

How is a giant sperm ejaculator formed? Development of the Zenker organ after the last moult in Pseudocandona marchica (Crustacea,Ostracoda, Candonidae)

‘Giant sperm’, in terms of exceptionally long spermatozoa, occur in various taxa in the animal kingdom, particularly in arthropods. In freshwater ostracods (Cypridoidea) reproducing with giant sperm, males are equipped with a pair of specialized ejaculators, or Zenker organs, located in the distal part of vas deferens.