Species‐specific habitat use of wing‐moulting waterbirds in response to temporary flightlessness

The choice of the moulting habitat is of paramount importance for wing‐moulting waterbirds that have to cope with a flightless period of several weeks. However, some species might have more restricted habitat requirements during moult than others, for example due to a highly specialized feeding ecology. The moult‐related habitat use of five species (Gadwall Anas strepera, Red‐crested Pochard Netta rufina, Common Pochard Aythya ferina, Tufted Duck Aythya fuligula, Coot Fulica atra) was compared at a European inland moulting site that offered a variety of water bodies characterized by different levels of nutrient concentration, water depth, shoreline vegetation density and disturbance. To determine location‐ and species‐specific densities, birds were regularly counted throughout the moulting seasons of 2010 and 2011. In 2011, additional data on Gadwalls were used to assess differences in requirements between the flightless phase of moult and the periods before and after. Furthermore, habitat choice of 38 tagged Gadwalls was compared among two to four successive years. During the moulting season, all species showed clear preferences for specific levels of nutrient content, suggesting an active choice of suitable food sources in both food specialists and generalists. Species showing the strongest attachment to shallow water (Gadwall and Coot) were most sensitive to human disturbance and increasing water depths, and species averse to diving (Gadwall) used ponds with dense shore vegetation while flightless. For Gadwalls, habitat conditions rather than nutrient supply became increasingly important during the flightless phase. Average return rates of 59 and 54% were recorded for male and female Gadwalls, respectively, and the repeated use of familiar locations could be demonstrated in the majority of returning birds (65%). Familiarity with the habitat apparently plays an important role and may enable individuals to compensate for suboptimal conditions at the moulting site.     

Testing the relationship between clutch size and brood size in the Coot (Fulica atra)

The time between egg laying and chick fledging is of crucial importance for the survival of young birds. I analyzed breeding output at consecutive phases of growth of young Coots (Fulica atra) relative to the clutch size and laying date. Considering the specific breeding biology of the Coot, I tested whether chick survival reveals clutch size-dependent variability. Clutch size did not affect hatching success; it only affected brood size, and that merely temporarily. During the first 20 days after hatching, i.e. during the time of the highest chick mortality, birds with larger clutches lost chicks at a higher rate. As a result, the number of fledged chicks was independent of the initial number of chicks, and pairs with different clutch sizes had a similar number of fledglings. The laying date had no effect. This pattern of age-related chick survival points to the greater role of the type of chick growth (semi-precocial) and behavior in their survival.     

Red-gartered Coot Fulica armillata feeding on the grapsid crab Cyrtograpsus angulatus: advantages and disadvantages of an unusual food resource

The behaviour of Red-gartered Coots feeding on an unusual food source was examined at Mar Chiquita Coastal Lagoon, Argentina. The grapsid crab Cyrtograpsus angulatus made up all observed prey items, and 61% were small. Both handling and foraging duration increased with the size of captured crabs, but foraging efficiency decreased. Crab availability affected both the dive duration of the Coots and their foraging decisions with regard to prey-size selection. Two species of gull were observed kleptoparasitizing Coots, especially when the Coot was handling medium or large crabs. Feeding by Coots on Cyrtograpsus angulatus has not been previously documented and may be a feeding innovation. Our estimations suggest that Coots were foraging optimally, since smaller crabs were more energetically profitable.     

Flightlessness affects cranial morphology in birds

Flightless birds belonging to phylogenetically distant clades share several morphological features in the pectoral and pelvic apparatus. There are indications that skull morphology is also influenced by flightlessness. In this study we used a large number of flightless species to test whether flightlessness in modern birds does indeed affect cranial morphology. Discriminant analyses and variation partitioning show evidence for a relationship between skull morphology and the flightless condition in birds. A possible explanation for the change in cranial morphology can be linked to the reduced selective force for light-weight skulls in flightless birds. This makes an increase in muscle mass, and therefore an enlargement of muscle insertion areas on the skull, possible. We also compared the ontogenetic trajectory of Gallus with the adult morphology of a sample of flightless species to see whether the apomorphic features characterizing the skull of flightless birds share the same developmental basis, which would indicate convergent evolution by parallelism. Skull morphology (expressed as principal component scores) of palaeognathous flightless birds (ratites) is dissimilar (higher scores) to juvenile stages of the chicken and therefore seem peramorphic (overdeveloped). Principal component scores of adult neognathous flightless birds fall within the range of chicken development, so no clear conclusions about the ontogenetic trajectories leading to their sturdier skull morphology could be drawn.     

Control of Aphis gossypii and Myzus persicae (Hemiptera: Aphididae) by a flightless strain of Harmonia axyridis (Coleoptera: Coccinellidae) on green pepper plants in open fields

The cotton aphid Aphis gossypii Glover and the green peach aphid Myzus persicae (Sulzer) are economically important pests with a worldwide distribution. We evaluated the efficacy of releasing a flightless adult strain of the multicolored Asian lady beetle Harmonia axyridis (Pallas) as a control measure against these aphids on green pepper plants in open fields. Flightless H. axyridis adults were observed on the green pepper plants in the releasing plots throughout the experimental period and were found to be effective biocontrol agents, markedly decreasing the numbers of aphids. These results suggest that adults of this flightless strain of H. axyridis are effective in controlling aphids on green pepper plants in open fields.     

Changes in abundance of water birds species in southern Bohemia (Czech Republic) in the last 10 years

In 1981–1992, the breeding fauna of 153 fishponds was studies in three fishpond regions in south Bohemia (Czechoslovakia). On each pond, all water and wetland birds were censused by the Two-check method, which consists of adult birds counts during each breeding season (the first in the second part of May, and the second in the second part of June). We assessed the numbers of all species living in water and wetland habitats in the breeding season.The decline in abundances (expressed as number of individuals) of ducks (Anatinae), grebes (Podicipediformes), Coot (Fulica atra), Moorhen (Galinulla chloropus), Black-headed Gull (Larus ridibundus) were the most marked changes recorded. Marked increase in abundances was found in Cormorant (Phalacrocorax carbo) and Mute Swan (Cygnus olor) during the whole decade, whereas in Graylag Goose (Anser anser) the increase was restricted to the first half of the period under study. Numbers of charadriform and passeriform birds fluctuated in the dependence of water level variation.     

A refined model of body mass and population density in flightless birds reconciles extreme bimodal population estimates for extinct moa

Flightless birds were once the largest and heaviest terrestrial fauna on many archipelagos around the world. Robust approaches for estimating their population parameters are essential for understanding prehistoric insular ecosystems and extinction processes. Body mass and population density are negatively related for extant flightless bird species, providing a method for quantifying densities and population sizes of extinct flightless species. Here we assemble an updated global data set of body mass and population densities for extant flightless birds and estimate the relationship between these variables. We use generalised least squares models that account for phylogenetic relatedness and incorporate the effects of limiting factors (e.g. habitat suitability) on population density. We demonstrate the applicability of this allometric relationship to extinct species by estimating densities for each of the nine species of moa (Dinornithiformes) and generating a combined spatially explicit map of total moa density across New Zealand. To compare our density estimates with those previously published, we summed individual species' abundances to generate a mean national density of 2.02–9.66 birds km⁻² for low- and high-density scenarios, respectively. Our results reconcile the extreme bimodality of previous estimates (< 2 birds km⁻² and > 10 birds km⁻²) and are comparable to contemporary densities of large herbivorous wild mammals introduced into New Zealand about 150 yr ago. The revised moa density has little effect on the harvest rates required to bring about extinction within 150–200 yr, indicating that rapid extinction was an inevitable response to human hunting, irrespective of the initial population of moa.     

Suppression of aphids by augmentative release of larvae of flightless Harmonia axyridis

The release of the larvae of flightless ladybird beetles could extend the duration of effective control because of the longer presence of adult stages, and the per individual production cost for juvenile stages is lower than that for adult stages. A preliminary release experiment was conducted to assess the effectiveness in controlling two aphid species, Aphis gossypii and Aulacorthum solani, using second instars of a flightless strain of Harmonia axyridis. The number of A. gossypii was suppressed in greenhouses that contained the flightless strain compared with greenhouses that contained the wild‐type strain. In one of two replicates, more flightless adults were observed on the plants for longer than wild‐type adults. However, it was not clear whether the extension of the residence period contributed to the effectiveness in controlling aphids directly, because the population of ladybirds in the greenhouses consisted of both larvae and adults during parts of the experimental period. In flightless H. axyridis, the release of larvae was more effective in suppressing A. solani than the release of adults. These results suggest that it may be more effective to release larvae of flightless H. axyridis than wild‐type larvae or flightless adults.     

Greater energy stores enable flightless moulting geese to increase resting behaviour

Many species of waterfowl undergo a post‐breeding simultaneous flight feather moult (wing moult) which renders them flightless and vulnerable to predation for up to 4 weeks. Here we present an analysis of the correlations between individual time‐budgets and body mass states in 13 captive Barnacle Geese Branta leucopsis throughout an entire wing moult. The daily percentage of time spent resting was positively correlated with initial body mass at the start of wing moult. Behaviour of individual birds during wing moult is dependent on initial physiological state, which may in turn be dependent on foraging ability; the storage of energy before the start of wing moult will help birds to reduce exposure to the dangers of predation.     

SOME NOTES ON THE BIRDS OF THE ISOKA DISTRICT OF THE NORTHERN PROVINCE OF NORTHERN RHODESIA

Dean, W. R. J. 1978. Moult seasons of some Anatidae in the western Transvaal. Ostrich 49:76-84.

Spurwinged Geese Plectropterus gambensis, Egyptian Geese Alopochen aegyptiacus, Yellow-billed Ducks Anas undulata, Redbilled Teal A. erythrorhyncha and Southern Pochard Netta erythrophthalma have a flightless moult mainly during the dry season, from April to August, in the western Transvaal. South African Shelduck Tadorna cana moult during October to February after breeding during July and August. The Cape Shoveller Anas smithii has two main flightless periods, April-May and October-January. Cape Teal A. capensis have been recorded in flightless moult in October, December and January.

The duration of the flightless period correlates with wing length; larger and longer winged Anatidae require proportionally more time for wing moult than do smaller and shorter winged Anatidae.

Geese and shelducks moult on large open lakes with an open shore. Ducks have been recorded flightless on lakes and dams, with or without emergent vegetation.     

Can birds count eggs in their nests?

Sensory mechanisms controlling avian clutch size have diversified into distinct types, according to the nature of the input that is used to disrupt the growth of ovarian follicles and hence halt egg‐laying. In an article on brood parasitism, Lyon (2003) claimed that female American Coots Fulica americana can reduce their clutch size on the basis of visual cues in response to eggs laid in their nests by other females; in this species, therefore, egg counting would be used to control clutch size. After a close examination of the physiological determination of clutch size in American Coots, I show that seven of 17 parasitized clutches were smaller than the range controlled through the mechanism using an input to disrupt follicular growth (7–10 eggs per clutch). My reanalysis suggests that American Coots are incapable of adjusting clutch size via counting and re‐asserts that a species that can count eggs has yet to be found among birds that rely upon their own body heat for incubation.     

The introduced clam Ensis americanus in the Wadden Sea: field experiment on impact of bird predation and tidal level on survival and growth

In the Danish Wadden Sea the intertidal distribution of the introduced bivalve Ensis americanus (syn. E. directus) is restricted to a narrow zone around the mean low water level. To test the possible impact of birds and submersion time on dynamics and distribution of the clams, adult specimens of E. americanus collected near the low water line were transplanted to two intertidal sites and established in open and net-covered experimental plots for 9 weeks (autumn 2001). The lowest survival of clams was registered at the low-shore-site (LSS) in plots open to bird predators, suggesting that birds such as Common Eider (Somateria mollissima) or Oystercatcher (Haematopus ostralegus) may control the abundance of E. americanus at the lower tidal levels. For clams showing increment in shell length during the study period, the shell growth rates were highest at the LSS and lowest in the open plots at the high-shore-site (HSS). Differences in immersion time and thus food supply may explain this pattern. Body mass index (BMI) of the clams showed basically the same pattern as the survivorship: lowest BMI in open plots at the LSS and highest in the covered plots at this site. Clams from the HSS were intermediate in their BMI. Disturbance by birds in the open plots at the LSS may explain the low BMI. In conclusion birds may be an important factor controlling abundance of E. americanus in the lower intertidal zone.     

The flightless period of free-living male Teal Anas crecca in northern Sweden

A total of 126 flightless adult male Teal was caught in 1982 and 1983 in a moulting area in northern Sweden (63° 53'N). The birds were weighed and the stage of development of their primaries and secondaries measured. During the flightless period, the weight of the birds decreased by an average of 10% in 1982 and 19% in 1983. This weight decrease allows the birds to fly before their primaries and secondaries are full-grown. The weight loss is considered to be an adaptation making it possible, for example, for lighter birds to escape predators sooner after moulting. Birds varied greatly in the dates at which they initiated their flightless period, that is, from the end of June to the end of July. Thus, it is probable that both local birds and birds from distant breeding grounds were present in the moulting area.     

Natural flightless morphs of the ladybird beetle Adalia bipunctata improve biological control of aphids on single plants

The challenge of using ladybird beetles for biological control of insect pests such as aphids is that the adult beetles tend to fly away from the host plants. Therefore, flightless ladybirds might improve biocontrol. There are several artificial ways to obtain flightless beetles, but it may be preferable to use natural variation in flight ability. We investigated, for the first time, biocontrol by inundative augmentation of natural flightless morphs of the ladybird beetle Adalia bipunctata. Microcosm experiments using single leaves with one of three species of aphid revealed no differences in consumption behavior between flightless and winged beetles. Monitoring for 48 h of single, caged pepper plants infested with aphids of Myzus persicae nicotianae or Aulacorthum solani showed that flightless beetles had a longer residence time on the plants than winged beetles. This only translated into significantly better biocontrol of M. persicae. Despite their difference in residence time, both beetle morphs reduced the population growth of A. solani. This is probably explained by the tendency of A. solani to drop from the plant upon disturbance, and we predict that flightless beetles may outperform winged ones in the long term. Overall, our results provide a proof of principle that natural flightless A. bipunctata can improve biocontrol of aphids by ladybird beetles. However, we recognize that the effect of biocontrol will vary with the species of aphid used and that further examination in long term and large scale experiments is required.     

Long-term trends,seasonal abundance and energy consumption of waterbirds at Rietvlei,Western Cape,South Africa, 1950–1997

Waterbird counts at Rietvlei, a freshwater wetland in the Western Cape, were analysed for four periods: 1950–56, 1961–67, 1980–85 and 1992–97. Ninety-eight species of waterbirds were recorded during these counts. Palaearctic migrants predominated during summer, with Curlew Sandpipers, Calidris ferruginea, being the mast abundant species in all years, except in the 1960s and 1950s when Red-knobbed Coots, Fulica cristata, and Ruffs, Philomachus pugnax, outnumbered them respectively. Red-knobbed Coots and Yellow-billed Ducks, Anas undulata, were the most numerous species during winter in all years. The numbers of waterbirds at Rietvlei have shown a progressive increase since the 1950s, reaching over 8300 birds in the mid-summer of 1997. The increase in abundance of several species is attributed to changes in the water regime and habitat availability at the wetland. In terms of waterbird abundance, Rietvlei is the most important temporary vlei in the Western Cape and it ranks 16th in terms of waterbird abundance compared with all South African coastal and inland wetlands for which counts are available. In the light of Rietvlei being proclaimed a nature reserve, a conservation management plan for the area is discussed.     

Flightlessness in the Galápagos cormorant (Compsohalieus [Nannopterum] harrisi): heterochrony, giantism and specialization

A study of flightlessness in the Galápagos cormorant (Compsohalieus [Nannopterum] harrisi) was undertaken using study skins and skeletons of C. harrisi and eight flighted confamilials; in addition, four skin specimens and disassociated skeletal elements of the extinct spectacled cormorant (C. perspicillatus) of Beringia, reputed by some to have been flightless, were studied. Anatomical specimens of C. penicillatus and C. harrisi were dissected for myological comparisons. Flightless C. harrisi is 1.6 to 2.2 times as heavy as its extant flighted congeners; males averaged 3958 g and females averaged 2715 g in total body weight. Estimates of body weight for C. perspicillatus based on femur length approximated 3900 g. Wing lengths of C. harrisi were smaller than those of any other cormorant, averaging 190 mm and 170 mm for males and females, respectively. Wing-loadings (g body mass.cm^-2 wing area) of flighted cormorants ranged from 1.0 to 1.7. Estimated wing-loadings, incorporating approximate wing areas, were 2.0 and 5.1 g.cm^-2 for C. perspicillatus and C. harrisi, respectively; the former suggests that C. perspicillatus was probably capable of laboured flight. The small wings of C. harrisi result from an c. 50% shortening of remiges, accompanied by reduced asymmetry of vane widths and increased rounding of the tips, and significant reductions in lengths of wing bones, particularly the radius and ulna. Numbers of primary and secondary remiges in C. harrisi remain unchanged. Multivariate morphometries revealed that sexual dimorphism in external and skeletal dimensions is significantly greater in C. harrisi than in flighted cormorants. Canonical analysis of six external measurements indicated that C. harrisi is distinguished primarily by its relatively short wings. Skeletal peculiarities of C. harrisi were diverse, including conformational changes in the sternum, furcula, coracoid, humerus, ulna, radius, carpometacarpus and patella. Mensural comparisons confirmed substantial reductions in elements of the pectoral girdle of C. harrisi, particularly the sternal carina, as well as the alar skeleton, especially the radius and ulna. Differential shortening of the wing elements resulted in significant differences in proportions within the wing skeleton. These unique skeletal proportions of C. harrisi, in addition to its great overall size, combine to produce an immense multivariate skeletal distance between C. harrisi and all confamilials. Sexual dimorphism in skeletal dimensions, in both total and size-corrected data, was 2–3 times greater in C. harrisi than in other phalacrocoracids sampled. Most pectoral muscles of C. harrisi were absolutely or relatively smaller than those of C. penicillatus, in spite of its larger body size. No muscles or parts thereof were lacking in the pectoral limb of C. harrisi, but a number of qualitative differences distinguished the musculature of the flightless species, including: an exceptionally tough skin involving a well-developed M. pectoralis pars abdominalis and M. latissimus dorsi interscapularis; a thin, medially obsolete and laterally extensive M. pectoralis pars thoracica; a weakly developed M. rhomboideus profundus consisting of a variably tendinous fascia invested with three fasciculi of muscle fibres; an extraordinarily thick, extensive M. obliquus externus abdominis, which, together with a unique cnemio-costal slip of smooth muscle, restricts the metapatagium through an anchoring of M. serratus superficialis metapatagialis; and the presence of a unique alular muscle named here as M. levator alulae. Fusions of the tendons of origin and insertion, respectively, of M. flexor digiti superficialis and M. flexor digiti profundus in C. harrisi, muscles derived from a common muscle primordium, and the retention of a carpometacarpal tendon of M. flexor carpi ulnaris cranialis constitute strong evidence of pectoral paedomorphosis in C. harrisi. Mensural comparisons quantified the reduction of pectoral muscles in C. harrisi and indicated that these reductions were especially pronounced in the distal musculature. Morphological characteristics of Phalacrocoracidae, together with the exploitation of localized marine food resources and weakly developed seasonal movements of Compsohalieus, may have predisposed the founding population of C. harrisi to flightlessness. Anatomical changes in C. harrisi are exceeded in degree among foot-propelled diving birds by those of only a few fossil flightless birds (e.g. Hesperomis, Chendytes). Many of the morphological peculiarities of C. harrisi are paedomorphic, although several are not attributable to developmental heterochrony. These morphological characters of flightless C. harrisi are considered with respect to locomotion, feeding ecology, reproduction and demography of the species, and are compared with those of other flightless carinates.     

Do Seaducks Minimise the Flightless Period?: Inter- and Intra-Specific Comparisons of Remigial Moult

Remigial moult is one of the crucial events in the annual life cycle of waterfowl as it is energetically costly, lasts several weeks, and is a period of high vulnerability due to flightlessness. In waterfowl, remigial moult can be considered as an energy-predation trade-off, meaning that heavier individuals would minimise the flightless period by increasing feather growth rate and energy expenditure. Alternatively, they could reduce body mass at the end of this period, thereby reducing wing-loading to increase flight capability. We studied timing of remigial moult, primary growth rates, flightlessness duration, and the pattern of body mass variation in 5 species of captive seaducks (Melanitta fusca, M. perspicillata, Clangula hyemalis, Histrionicus histrionicus, and Somateria mollissima) ranging in size from 0.5 to 2.0 kg. Their feather growth rates weakly increased with body mass (M^0.059) and no correlation was found at the intra-specific level. Consequently, heavier seaduck species and especially heavier individuals had a longer flightless period. Although birds had access to food ad libidum, body mass first increased then decreased, the latter coinciding with maximum feather growth rate. Level of body mass when birds regained flight ability was similar to level observed at the beginning of remigial moult, suggesting they were not using a strategic reduction of body mass to reduce the flightlessness duration. We suggest that the moulting strategy of seaducks may be the result of a compromise between using an intense moult strategy (simultaneous moult) and a low feather growth rate without prejudice to feather quality. Despite the controlled captive status of the studied seaducks, all five species as well as both sexes within each species showed timing of moult reflecting that of wild birds, suggesting there is a genetic component acting to shape moult timing within wild birds.     

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.     

Geese and Red-knobbed Coot on the Kafue Flats in Zambia, 1970–1974

The occurrence, chronology of breeding and wing-moult, and feeding habits of the Pigmy Goose Nettapus auritus, Knob-billed Goose Sarkidiornis melanotos, Spur-winged Goose Plectropterus gambensis, Egyptian Goose Alopochen aegyptiaca and Red-knobbed Coot Fulica cristata on the Kafue Flats in Zambia are described based on observations made between October 1970 and January 1974 at Lochinvar National Park and, from the air, over the central section of the Flats. The Pigmy Goose, Knob-billed Goose, Egyptian Goose and Red-knobbed Coot fed largely by grazing. They were most abundant between March and June, occurring in greatest numbers at Lochinvar. Egyptian Geese occasionally nested on the flood-plain at Lochinvar between August and January, but there was no proof of the other species nesting there. Many Pigmy Geese and male Knob-billed Geese moulted on the lagoon at Lochinvar between March and June; small numbers of Egyptian Geese and Red-knobbed Coot also moulted in the Park, the former between February and May, the latter in June and July. The Spur-winged Goose fed mostly by grazing and grubbing. Numbers on the Flats were greatest between June and November when 60,000–90,000 moved downriver as the floods subsided. Many bred between January and March in the fringing zone and moulted on the floodplain between May and July. A hydroelectric scheme, completed in 1975, may benefit the Pigmy Goose and Red-knobbed Coot but harm the Spur-winged Goose.     

The legs of ostriches (Struthio) and moas (Pachyornis)

Ostriches were filmed running at maximum speed, and forces on the feet were calculated. Measurements were made of the principal structures in the legs of an ostrich. Hence peak stresses in muscles, tendons and bones were calculated. They lay within the range of stresses calculated for strenuous activities of other vertebrates. The ostrich makes substantial savings of energy in running, by elastic storage in stretched tendons. Pachyornis was a flightless bird, much heavier than ostriches and with massively thick leg bones. These bones are shorter than predicted for its estimated body mass, by extrapolation from allometric equations for flying birds. An attempt is made to calculate the stresses that acted in the leg bones in running, for all possible patterns of leg movement. The stresses were probably rather low, unless Pachyornis was capable of running fast. It is argued that the optimum factor of safety for moo leg bones may have been exceptionally high, as a consequence of the absence of predators.