The Long and the Short of It: No Dietary Specialisation between Male and Female Western Sandpipers Despite Strong Bill Size Dimorphism

Many bird species show spatial or habitat segregation of the sexes during the non-breeding season. One potential ecological explanation is that differences in bill morphology favour foraging niche specialisation and segregation. Western sandpipers Calidris mauri have pronounced bill size dimorphism, with female bills averaging 15% longer than those of males. The sexes differ in foraging behaviour and exhibit partial latitudinal segregation during the non-breeding season, with males predominant in the north and females in the south. Niche specialisation at a local scale might account for this broad geographic pattern, and we investigated whether longer-billed females and shorter-billed males occupy different foraging niches at 16 sites across the non-breeding range. We used stable-nitrogen (δ¹⁵N) isotope analysis of whole blood to test for dietary specialisation according to bill length and sex. Stable-nitrogen isotope ratios increase with trophic level. We predicted that δ¹⁵N values would increase with bill length and would be higher for females, which use a greater proportion of foraging behaviour that targets higher-trophic level prey. We used stable-carbon (δ¹³C) isotope analysis to test for habitat segregation according to bill length and sex. Stable-carbon isotope ratios vary between marine- and freshwater-influenced habitats. We predicted that δ¹³C values would differ between males and females if the sexes segregate between habitat types. Using a model selection approach, we found little support for a relationship between δ¹⁵N and either bill length or sex. There was some indication, however, that more marine δ¹³C values occur with shorter bill lengths. Our findings provide little evidence that male and female western sandpipers exhibit dietary specialisation as a function of their bill size, but indicate that the sexes may segregate in different habitats according to bill length at some non-breeding sites. Potential ecological factors underlying habitat segregation between sexes include differences in preferred habitat type and predation risk.     

Parental investment asymmetries of a globally endangered scavenger: unravelling the role of gender,weather conditions and stage of the nesting cycle

ABSTRACT

Capsule: In Egyptian Vultures Neophron percnopterus, both sexes invested similar parental effort throughout the breeding period. However, there was variation in the degree of intensity of parental care during some stages of the breeding period, suggesting that sex-role specialization exists for some activities.

Aims: To quantify parental care behaviour of Egyptian Vultures for the first time and to examine the role of sex, weather conditions, and stage of nesting cycle on breeding ecology.

Methods: We monitored 15 nests of Egyptian Vultures to analyse parental care investment. We collected data on nine different behavioural parameters/activities per sex, which were recorded throughout the entire breeding period. Variation in parental investment was analysed using generalized linear mixed models.

Results: Females invested more effort in incubation/brooding (61.45% for females and 31.54% for males) and egg turning (0.45?events/h for females and 0.37?events/h for males) while males contributed more to nest material delivery to the nest (0.67?deliveries/h for males and 0.14?deliveries/h for females). Conversely, both sexes invested the same effort in nestling attendance (21.89% for females and 21.21% for males) and food provisioning (0.28?items/h for females and 0.25?items/h for males). Furthermore, parental investment was not affected by weather, especially during critical moments such as incubation/brooding, however, changeover rate was positively related to temperature.

Conclusion: Our results suggest that, in the Egyptian Vulture, one sex is not entirely responsible for a particular task and the compensatory effort of the other mate is required. Finally, our findings indicate that major events such as incubation onset and hatching caused important shifts in the patterns of parental investment.     

Sexing Wrens Troglodytes troglodytes indigenus using morphological measurements and discriminant analysis

Within the breeding population of a deciduous woodland, male Wrens averaged slightly larger than females in wing length, foot length, head plus bill length and body mass. First-year and older birds were similar for three of these morphometric variables, but the first-year group were shorter-winged than the older birds (by 3%). Despite this, wing length still proved to be a useful discriminator of sex when the data were pooled for all ages (74% of males exceeding the maximum female value of 49.8 mm). A discriminant function utilizing wing length and head plus bill length correctly sexed 96% of individuals in the sample from which it was derived (n = 49 males, 36 females). This function was converted into a predictive equation for the probability of an unknown case being male, and a plot of wing length against head plus bill length along with probability contours which enabled the sexing of unknown cases without the need for calculations. Measurements of the morphological variables investigated were found to be highly repeatable for a single worker, with the exception of body mass which differed substantially between the sexes but which was only moderately repeatable. The methods provided should prove valuable for sexing British mainland Troglodytes troglodytes indigenus, and probably also Western European T. t. troglodytes, but should not be applied to the larger, offshore island races.     

Sex-Based Differences in Adélie Penguin (Pygoscelis adeliae) Chick Growth Rates and Diet

Sexually size-dimorphic species must show some difference between the sexes in growth rate and/or length of growing period. Such differences in growth parameters can cause the sexes to be impacted by environmental variability in different ways, and understanding these differences allows a better understanding of patterns in productivity between individuals and populations. We investigated differences in growth rate and diet between male and female Adélie Penguin (Pygoscelis adeliae) chicks during two breeding seasons at Cape Crozier, Ross Island, Antarctica. Adélie Penguins are a slightly dimorphic species, with adult males averaging larger than adult females in mass (~11%) as well as bill (~8%) and flipper length (~3%). We measured mass and length of flipper, bill, tibiotarsus, and foot at 5-day intervals for 45 male and 40 female individually-marked chicks. Chick sex was molecularly determined from feathers. We used linear mixed effects models to estimate daily growth rate as a function of chick sex, while controlling for hatching order, brood size, year, and potential variation in breeding quality between pairs of parents. Accounting for season and hatching order, male chicks gained mass an average of 15.6 g d^-1 faster than females. Similarly, growth in bill length was faster for males, and the calculated bill size difference at fledging was similar to that observed in adults. There was no evidence for sex-based differences in growth of other morphological features. Adélie diet at Ross Island is composed almost entirely of two species—one krill (Euphausia crystallorophias) and one fish (Pleuragramma antarctica), with fish having a higher caloric value. Using isotopic analyses of feather samples, we also determined that male chicks were fed a higher proportion of fish than female chicks. The related differences in provisioning and growth rates of male and female offspring provides a greater understanding of the ways in which ecological factors may impact the two sexes differently.     

Sex determination of Antarctic Petrels Thalassoica antarctica by discriminant analysis of morphometric characters

We present data on sexual dimorphism in some morphological measurements (wing length, head length, bill depth and bill length) in the Antarctic Petrel Thalassoica antarctica. Males were on average larger than females for all measurements. Sexual dimorphism was on average largest for bill depths whereas wing lengths discriminated least between the sexes. A discriminant function including bill depth, head length and wing length correctly sexed 92% of the sample. Due to between-measurer variation it is recommended that morphometric measurements obtained by others on sexed birds are compared with ours before proceeding with the use of the discriminant function on unsexed individuals.Publication No. 116 of the Norwegian Antarctic Research Expeditions (1991/92)     

The haematozoan parasites of Common Blackbirds Turdus merula: associations with host condition

A population of Common Blackbirds Turdus merula was studied to investigate the relationships between the presence of blood parasites and host morphometrics, a putative sexually selected trait (bill colour), and reproductive parameters. Haematozoa of four genera were detected and their prevalence was high. Infection was negatively associated with adult morphometrics: adults infected with Leucocytozoon were in relatively poor body condition and had shorter wings than uninfected birds. The bill colour of males infected with Plasmodium tended to be duller than that of uninfected males, and in females Haemoproteus infection was significantly positively associated with bill coloration. Haematozoan infection of females was unrelated to measured reproductive parameters, and there was no relationship between blood parasite infection and the provision of parental care.     

Sex determination of African Penguins Spheniscus demersus using bill measurements: method comparisons and implications for use

African Penguins Spheniscus demersus are sexually dimorphic; on average, males are larger than females but measurements overlap making sex determination difficult through observations alone. We developed a discriminant function, using bill length and depth from a sample of birds sexed from gonad visualisation during post-mortem, which correctly classified 93% of the individuals. Cross-validation correctly assigned 90% of DNA-sexed birds and 91% of birds sexed by partner measurement comparisons. The use of discriminant function score cutpoints, while leaving 16% and 29% of birds unclassified, improved accuracy of birds sexed by DNA to 97% and of those sexed by partner comparison to 99%. Bill depth was found to be a discriminating variable. However, two techniques for measuring bill depth are currently in use for African Penguins. While these measurements are correlated (r = 0.85), they differ on average by 1.4?mm hindering accuracy of sex determination when using a discriminant function developed from the other bill depth measurement. Exploration of adult bill morphology of birds sexed from DNA at different colonies suggests the discriminant functions can be applied throughout the African Penguins’ South African range.     

Sex differences in survival selection in the serin,Serinus serinus

Natural selection is demonstrated in most natural populations which suggests that populations are dispatched from their adaptive peaks as a result of selection on correlated characters, or conflicting selection between the sexes. We analysed patterns of survival selection in a population of serins (Serinus serinus) outside Barcelona over a period of 13 years. There was directional selection for increased wing length in males and females accompanied by strong disruptive selection on both tail and wing length in males and a selection against a positive correlation between the two characters in males. In females there was directional selection for increased bill width but decreased bill depth, which should be contrasted to the stabilizing selection acting on bill depth in males. There were conflicting selection on the characters within a sex and conflicting selection of the same characters between sexes, which constrain the rate of access to the nearest adaptive peak.     

基于形态特征判定五种鹬类性别的可靠性

快速准确地鉴定两性同型鸟类个体性别在鸟类生态学研究中具有重要意义。本文选择2008年春季迁徙期在崇明东滩停歇的大滨鹬(Calidris tenuirostris)、红腹滨鹬(C.canutus)、红颈滨鹬(C.ruficollis)、尖尾滨鹬(C.acuminata)及翘嘴鹬(Xenus cinereus)5种两性同型的鹬类,用分子生物学方法进行性别鉴定,并基于个体的形态特征(体重、翅长、喙长、头喙长及跗跖长)采用判别分析方法对性别进行判定。结果表明,尖尾滨鹬雄性各形态特征均显著大于雌性,其他4种鹬类则相反。5种鹬类形态特征的性别差异指数在0.5%~25.3%之间,重叠度在29.4%~98.6%之间。5种鹬类判别分析判定性别的准确率在(0.69±0.06)~(0.96±0.01)之间,其中,尖尾滨鹬判别准确率(0.96)最高,翘嘴鹬判别准确率(0.69)最低。形态特征在两性间的差异程度影响性别的判别准确率。另外,两性性比对性别判别的准确率也有影响:性比偏雄性鸟类的雄性判别准确率高于雌性,而性比偏雌性鸟类的雌性判别准确率高于雄性。采用判别分析估测的性比与分子生物学鉴定结果相似,表明判别分析在判定种群的性比方面具有较高的可靠性。     

Sexual size dimorphism and assortative mating in Carolina Wrens

ABSTRACT.   Sexual size dimorphism (SSD) may be due to sexual and natural selection, but identifying specific mechanisms that generate such dimorphism in a species is difficult. I examined SSD in Carolina Wrens ( Thryothorus ludovicianus ) by examining (1) the degree of SSD in the population and between pairs using five morphometrics, (2) assortative mating patterns based on size and age, and (3) relationships between size and longevity. Analysis revealed that males were significantly larger than females in all body measurements. For example, mass, bill, and wing measurements yielded a canonical variable that permitted separation of the sexes and linear classification functions correctly determined the sex of 95% (238/250) of all wrens measured. No evidence was found to suggest that SSD was related to resource partitioning. However, assortative mating trends based on morphometrics (e.g., wing length), positive associations between longevity and morphometrics (e.g., wing length in females and body size in males), and intense male-male contests for territorial resources year-round provide evidence that sexual selection may contribute to SSD in Carolina Wrens.     

Sex‐specific winter distribution in a sexually dimorphic shorebird is explained by resource partitioning

Sexual size dimorphism (SSD) implies correlated differences in energetic requirements and feeding opportunities, such that sexes will face different trade‐offs in habitat selection. In seasonal migrants, this could result in a differential spatial distribution across the wintering range. To identify the ecological causes of sexual spatial segregation, we studied a sexually dimorphic shorebird, the bar‐tailed godwit Limosa lapponica, in which females have a larger body and a longer bill than males. With respect to the trade‐offs that these migratory shorebirds experience in their choice of wintering area, northern and colder wintering sites have the benefit of being closer to the Arctic breeding grounds. According to Bergmann's rule, the larger females should incur lower energetic costs per unit of body mass over males, helping them to winter in the cold. However, as the sexes have rather different bill lengths, differences in sex‐specific wintering sites could also be due to the vertical distribution of their buried prey, that is, resource partitioning. Here, in a comparison between six main intertidal wintering areas across the entire winter range of the lapponica subspecies in northwest Europe, we show that the percentage of females between sites was not correlated with the cost of wintering, but was positively correlated with the biomass in the bottom layer and negatively with the biomass in the top layer. We conclude that resource partitioning, rather than relative expenditure advantages, best explains the differential spatial distribution of male and female bar‐tailed godwits across northwest Europe.     

Size,shape and sex differences in three subspecies of Black-tailed Godwits Limosa limosa

ABSTRACT

Capsule: Black-tailed Godwits Limosa limosa show sexual size dimorphism and size differences between the subspecies. The shape varies slightly between the subspecies, but not between the sexes.

Aims: To investigate whether and how the three subspecies of Black-tailed Godwits, and the sexes of these subspecies, differ in size and shape.

Methods: We collected body dimensions (lengths of the bill, total head, tarsus, tarsus-toe and wing) of adult Black-tailed Godwits from three locations (Iceland, the Netherlands and northwest Australia) corresponding to the breeding or wintering grounds of three known subspecies (islandica, limosa and melanuroides, respectively). Determining sex by molecular assays, we computed degrees of sexual size dimorphism. Using principal component analysis (PCA), we compared differences in size and shape among the different subspecies.

Results: The limosa subspecies was the largest and also showed the most significant sexual size dimorphism. Sexual size dimorphism was smallest for wing length and largest for bill length. The first two axes of the PCA that included all subspecies of both sexes explained 94% of the total variation. Most body dimensions were highly correlated with each other, but wing length varied independently of the other dimensions. Males and females differed only in size (the first axis). However, one of the two small subspecies, islandica, also differed in shape (the second axis) from limosa and melanuroides.

Conclusions: In all three subspecies of Black-tailed Godwits, females are larger than males. The fact that subspecies differed in the degree of size dimorphism and slightly in shape hints at sex-related differences in the ecological selection pressures between the different flyways.     

ON THE SOUTH AFRICAN RACE OF THE LITTLE SPOTTED WOODPECKER

Discriminant analysis functions have previously been determined for sexing Mediterranean Yellow-legged Gulls Larus michahellis michahellis from the western Mediterranean basin. However, data from eastern Mediterranean populations are lacking. In this work, we used morphometric data from a sample of 81 Mediterranean Yellow-legged Gulls (39 males and 42 females) breeding in the Gulf of Gabès in south-eastern Tunisia to (1) determine a discriminant function useful for sex discrimination, and (2) assess the accuracy of previously published functions in sexing Mediterranean Yellow-legged Gulls from our study area. Our results showed marked sexual differences in all morphological measurements, with males being significantly larger than females. The best discriminant function included head length, bill depth and wing length, and accurately classified 93% of sampled birds. We also found that Mediterranean Yellow-legged Gulls from the Gulf of Gabès could accurately be sexed using discriminant functions determined for another North African population, but not with a function determined for a South European population, although distances between sites are almost the same.     

Variance‐Sensitive Green Woodhoopoes: A New Explanation for Sex Differences in Foraging?

Studies of cooperatively breeding birds rarely benefit from the extensive research on adaptive foraging behaviour, despite the potential for concepts such as state‐dependent foraging to explain many aspects of behaviour in social groups. For example, sex differences in preferred foraging techniques used by green woodhoopoes, Phoeniculus purpureus, have previously been explained by sexual dimorphism in bill length and the benefits afforded by foraging specialization and niche differentiation within cooperative groups. Contrary to this argument, there were no sex differences in mean foraging success and/or prey size captured when males and females used the same foraging techniques. Subordinates of both sexes did experience lower and more varied foraging success compared with dominants, but probably only as a consequence of competition or inexperience. However, dominant males experienced greater variance in individual foraging success compared with dominant females, and dominant males also experienced greater variances in prey size when using their preferred foraging techniques. Dominant males therefore appeared to specialize in foraging techniques that provided more variable rewards, whilst dominant females consistently chose to minimize variation in reward. Dominant females also experienced less variance in foraging returns when using the same techniques as males, suggesting a possible link with sexual dimorphism in bill length. Partitioning of foraging niches in dominant green woodhoopoes therefore appears to be better explained by sex differences in variance (risk) sensitivity to foraging rewards. We suggest that this kind of detailed analysis of state‐dependent foraging has the potential to explain many of the crucial age and sex differences in behaviour within cooperative groups.     

White‐headed Vulture Trigonoceps occipitalis shows visual field characteristics of hunting raptors

The visual fields of the Aegypiinae vultures have been shown to be adapted primarily to meet two key perceptual challenges of their obligate carrion‐feeding behaviour: scanning the ground and preventing the sun's image falling upon the retina. However, field observations have shown that foraging White‐headed Vultures Trigonoceps occipitalis are not exclusively carrion‐feeders; they are also facultative predators of live prey. Such feeding is likely to present perceptual challenges that are additional to those posed by carrion‐feeding. Binocularity is the key component of all visual fields and in birds it is thought to function primarily in the accurate placement and time of contact of the talons and bill, especially in the location and seizure of food items. We determined visual fields in White‐headed Vultures and compared them with those of two species of carrion‐eating Gyps vultures. The visual field of White‐headed Vultures has more similarities with those of predatory raptors (e.g. accipitrid hawks) than with the taxonomically more closely related Gyps vultures. Maximum binocular field width in White‐headed Vultures (30°) is significantly wider than that in Gyps vultures (20°). The broader binocular fields in White‐headed Vultures probably facilitate accurate placement and timing of the talons when capturing evasive live prey.     

A STUDY OF THE CINNAMON-BREASTED ROCK BUNTING Fringillaria tahapisi tehepisi

Brown, C. J. &; Piper, S. E. 1988. Status of Cape Vultures in the Natal Drakensberg and their cliff site selection. Ostrich 59:126-136.

Ground, aerial and questionnaire surveys on the status of the Cape Vulture Gyps coprotheres in the Little and High Drakensberg mountain ranges of Natal were carried out from 1981–1983. The area supported at least 1325 Cape Vultures, 60% on the High Drakensberg in 17 nesting colonies (mean of 31 birds per colony) and 38 roosts (mean of 7 birds). In the Little Drakensberg 17 sites were found, six of which were confirmed nesting colonies (mean of 53 birds) and seven were roosts (mean of 16 birds). A minimum of 215 nests was recorded while the actual number of breeding pairs was probably about 325. In the Natal Drakensberg 84% of nests and 77% of roosting birds occupied ledges that faced between east and south, on the lee side of the generally westerly and northwesterly “Bergwind”. Nesting ledges in the High Drakensberg were on average about 100 m lower than roosting ledges. The northern Drakensberg supported the largest numbers of Cape Vultures (19 birds per 10 km of cliffs), while the smallest numbers were recorded in the south (8 birds per 10 km of cliffs). In the south smallstock farming predominates and the availability of carrion is higher than in the north where mainly largestock are farmed. The use of poisons (mainly strychnine) for killing mammalian predators on farms is considered to be the main reason for fewer birds occurring in the south, and is the main threat to Cape Vultures in the Natal Drakensberg.     

Sexual size dimorphism in red-necked phalaropes and functional significance of nonsexual bill structure variation for feeding performance

Sexual size dimorphism is widespread in shorebirds, yet no tests of the assumption that such size dimorphism extends to functionally significant dimensions of the bill exist. This report presents tests of: (1) the assumption that sexual size dimorphism extends to the feeding structures in sexually size dimorphic bird, and (2) the hypothesis that bill-size variation influences feeding performance in Phalaropus lobatus, the red-necked phalarope. Discriminant function analysis revealed that the sexes of this species can be distinguished on the basis of five body size/bill length variables, but with low accuracy in sexing of females because of misclassification of small females as males. In the shorebird literature, the assumption is generally made that in the absence of selection to the contrary, bill size scales to body size and hence sexual size dimorphism extends to bill size. However, discriminant function analysis of measures from red-necked phalaropes failed to separate the sexes on the basis of either external or internal bill dimensions other than length. Nonetheless, internal dimensions of the upper jaw combined with exposed culmen length explained 86% of the variance in feeding performance of phalaropes; high feeding performance depends on a wide, shallow, complex internal bill structure. This study provides evidence that internal bill dimensions determine feeding performance in a manner consistent with the mechanics of surface tension transport of prey. These results suggest that some dimensions of bill size may be constrained by performance demands and demonstrate that variation in bill morphology has functional consequences. © 1996 Wiley-Liss, Inc.     

Sexual differences in head form and diet in a population of Mexican lance‐headed rattlesnakes,Crotalus polystictus

Sexual dimorphism of phenotypic traits associated with resource use is common in animals, and may result from niche divergence between sexes. Snakes have become widely used in studies of the ecological basis of sexual dimorphism because they are gape‐limited predators and their head morphology is likely to be a direct indicator of the size and shape of prey consumed. We examined sexual dimorphism of body size and head morphology, as well as sexual differences in diet, in a population of Mexican lance‐headed rattlesnakes, Crotalus polystictus, from the State of México, Mexico. The maximum snout–vent length of males was greater than that of females by 21%. Males had relatively larger heads, and differed from females in head shape after removing the effects of head size. In addition, male rattlesnakes showed positive allometry in head shape: head width was amplified, whereas snout length was truncated with increased head size. By contrast, our data did not provide clear evidence of allometry in head shape of females. Adults of both males and females ate predominately mice and voles; however, males also consumed a greater proportion of larger mammalian species, and fewer small prey species. The differences in diet correspond with dimorphism in head morphology, and provide evidence of intersexual niche divergence in the study population. However, because the sexes overlapped greatly in diet, we hypothesize that diet and head dimorphisms in C. polystictus are likely related to different selection pressures in each sex arising from pre‐existing body size differences rather than from character displacement for reducing intersexual competition. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 633–640.     

Parallel evolution of the sexes? Effects of predation and habitat features on the size and shape of wild guppies

Environmental gradients often lead to the parallel evolution of populations and species. To what extent do such gradients also lead to parallel evolution of the sexes? We used guppies (Poecilia reticulata) to examine the parallel and independent (sex‐specific) aspects of population divergence in response to predation and habitat features. Geometric morphometrics was used to analyse size and shape variation for 1335 guppies from 27 to 31 sites sampled in each of 2 years. Body size showed strong parallel population divergence; both sexes were larger at sites with a more open canopy and with higher flow. Body shape showed a mixture of parallel and independent population divergence. The strongest and most consistent effects were (1) high‐predation sites had males with smaller heads and deeper caudal peduncles, (2) open‐canopy sites had females with smaller heads and more distended abdomens and (3) high‐flow sites had males and females with smaller heads and deeper caudal peduncles.     

Lack of diet segregation during breeding by male and female Northern Flickers foraging on ants

Sexual size dimorphism can result in reduced competition if it leads males and females to use different foraging techniques or consume different prey items. Among woodpeckers, differences between males and females in bill length are common and may explain foraging differences in this family of birds. Northern Flickers (Colaptes auratus) are ground‐foraging woodpeckers that specialize on ants. However, the overall contribution of ants to their diet and the proportions of particular ant genera in their diet are not well known. To understand the relationship between bill morphology and the consumption of prey items, we compared the bill length and bill width of male and female flickers. We then collected and analyzed fecal samples from breeding flickers (N = 40 males, 33 females) at a study site in central British Columbia, Canada. Bills of male flickers were significantly longer (4%) and wider (5%) than those of females. Of 11 prey types identified, ants made up over 99% of their diet, and the abundance and composition of ant taxa in the diet did not differ between the sexes. We found significant year and time of season effects, with the abundance of Tapinoma sessile and Lasius spp. increasing from May to the end of June and differing between years. This difference in diet composition between years may have been due to changes in the abundance or accessibility of certain ant taxa related to differences in vegetation structure or weather. Nine ant taxa were consumed by flickers and the four most common were T. sessile, Lasius spp.,Myrmica spp., and the Formica fusca species group. The degree of dimorphism in bill size of male and female Northern Flickers in our study was smaller than reported for several species of arboreal‐foraging woodpeckers, suggesting that bill size of ground‐foraging woodpeckers may not be strongly linked to niche separation at the level of prey selection.