Intersexual niche segregation among three bark-foraging birds of eucalypt forests

All three species of specialized bark-climbing birds in eucalypt forests of north-eastern New South Wales exhibited sexual differences in their foraging behaviour. The degree of these differences was weakest in the least social species and strongest in the most social species, suggesting that intersexual niche segregation is an adaptation to reduce intraspecific competition for food. In the least social White-throated Treecreeper, the slightly longer-billed males foraged on dead branches more than females only during the breeding season when pair members were forced to share the same area. In the group-living Red-browed Treecreeper and Varied Sittella, males foraged lower and used rough-barked trunks and limbs more than females throughout the year. Sexual differences in the foraging substrates of Sittellas may have been related to sexual dimorphism of the bill, but in the monomorphic Red-browed Tree-creeper, male dominance was probably important. Some evidence suggests that further subdivision of the niche may occur in the highly social Sittella along the vertical dimension according to age and/or reproductive status. This report argues that intersexual niche segregation is just as likely to develop under conditions of increased interspecific competition as under reduced competition.     

Sexual dimorphism in relation to winter foraging in the white-backed woodpecker ( Dendrocopos leucotos)

Male white-backed woodpeckers (Dendrocopos leucotos) in a 250-km² study area in western Norway are significantly larger than females in bill length and depth, wing and tarsus lengths, and bodyweight. During the winters (October–March 1985–2002), most pairs were observed within their breeding territory where both sexes foraged mainly in grey alder and birch trees, and visited trees of the same tree height and stem width. However, males foraged more frequently on dead trees and on trees broken by storms. Males also used more trees with less bark cover, foraged nearer the ground and used foraging sites of larger diameter. Furthermore, males practised more deep wood-pecking and less bark-pecking than females. Unlike in other sexually dimorphic woodpecker species, the foraging niche breadth in wintering white-backed woodpeckers showed only minor sexual differences, and the sexes overlapped significantly in all parameters examined. Since previous studies in the area have shown that the sexes overlap considerably in use of their territory, it was expected, as found in other size dimorphic woodpeckers, that the larger male would displace the supposedly socially subordinate female to suboptimal feeding sites. In our area, the sexes were rarely seen together, and no sign of aggression between the sexes was observed. Despite the sex-specific differences found in the foraging behaviour of the birds, it is not obvious how the differences should be related to size dimorphism.Communicated by F. Bairlein     

The effect of social dominance on foraging by the Three-toed Woodpecker Picoides tridactylus

During the winter, female Three-toed Woodpeckers Picoides tridactylus , when unaccompanied by the male, foraged at a lower height above the ground and on tree trunks of greater diameter than when foraging together with a male. Among males, however, no such differences were found between those birds foraging in the absence of, or in company with, a female. The niche breadth of the males was less than that of the females, indicating a higher degree of specialization by the males. The niche overlap between the sexes was less when the birds foraged together in pairs, especially as regards foraging height and foraging site diameter. The intersexual segregation of foraging niche recorded for the Three-toed Woodpecker is therefore probably not determined genetically, but is due to social dominance by males which obliges the females to occupy the less preferred niche. Both sexes spent less time being vigilant when foraging together, than when alone.     

Ecological selection and sexual dimorphism in the sooty oystercatcher,Haematopus fuliginosus

Male and female sooty oystercatchers (subspecies Haematopus fuliginosus fuliginosus; Haematopodidae) have an average difference in bill length of 19%. We studied the relationship between this sexual dimorphism and foraging ecology at coastal sites in southern New South Wales, Australia. Intersexual foraging divergence was most striking in diet, with seven prey classes eaten exclusively by one sex (male: 4, female: 3), and all shared prey classes eaten in different proportions. Intersexual diet partitioning was also observed in energetic rewards gained from foraging, with females gaining highest energetic benefits from eating ascidians and males from eating limpets. Furthermore, within the most commonly consumed prey item, limpets, females gained higher energetic benefit from eating smaller sizes while males gained greater rewards from the largest limpet sizes. Intersexual divergence was also observed in several aspects of foraging behaviour. Finally, there was a significant effect of tidal cycles upon intersexual niche partitioning in this species; the degree of diet divergence varied between tide conditions and females had a consistently more efficient dietary intake on neap tides than males. Diet divergence in the sooty oystercatcher is greater than previously observed in any oystercatcher, and is correlated with the largest sexual bill dimorphism recorded in this family. It is argued that intersexual competition between territorial pairs is operating to diverge male and female bill morphology.     

Intersexual segregation in foraging microhabitat use by Magellanic Woodpeckers (Campephilus magellanicus): Seasonal and habitat effects at the world's southernmost forests

Animals facing seasonal food shortage and habitat degradation may adjust their foraging behaviour to reduce intraspecific competition. In the harsh environment of the world's southernmost forests in the Magellanic sub‐Antarctic ecoregion in Chile, we studied intersexual foraging differences in the largest South American woodpecker species, the Magellanic Woodpecker (Campephilus magellanicus). We assessed whether niche overlap between males and females decrease when food resources are less abundant or accessible, that is, during winter and in secondary forests, compared to summer and in old‐growth forests, respectively. We analysed 421 foraging microhabitat observations from six males and six females during 2011 and 2012. As predicted, the amount of niche overlap between males and females decreased during winter, when provisioning is more difficult. During winter, males and females (i) used trees with different diameter at breast height (DBH); (ii) fed in trunk sections with different diameters; and (iii) fed at different heights on tree trunks or branches. Vertical niche partitioning between sexes was found in both old‐growth and secondary forests. Such a niche partitioning during winter may be a seasonal strategy to avoid competition between sexes when prey resources are less abundant or accessible. Our results suggest that the conservation of this forest specialist, dimorphic and charismatic woodpecker species requires considering differences in habitat use between males and females.     

Sexual niche partitioning in two species of New Guinean Pachycephala whistlers

Sex differences in foraging behavior have been widely reported in the ornithological literature, but few examples are available from tropical avifaunas. Differences between males and females in foraging behavior have been hypothesized to be a byproduct of sexual size dimorphism or a result of niche partitioning to reduce intersexual competition for food or different reproductive roles. From 2010 to 2013, I used foraging data and mist‐net capture rates from multiple study sites to examine possible sex differences in the foraging behavior of two New Guinean Pachycephala whistlers. I found that male Regent (Pachycephala schlegelii) and Sclater's (Pachycephala soror) whistlers consistently foraged in higher strata than females. It is unlikely that these differences are due to sexual dimorphism because these species exhibit little sexual dimorphism. Sex differences in foraging behavior were consistent across years and study sites and did not appear linked to breeding behavior, supporting the food‐competition hypothesis, but not the reproductive‐roles hypothesis. Male territorial defense often occurs in relatively high strata in Pachycephala whistlers, possibly influencing male foraging strata. However, male territorial behavior cannot explain why females predominately forage in lower strata. Instead, intersexual competition for food resources is likely the primary driver of differences in the foraging behavior of male and female Regent and Sclater's whistlers.     

Intersexual trophic niche partitioning in an ant-eating spider (Araneae: Zodariidae)

Background

Divergence in trophic niche between the sexes may function to reduce competition between the sexes (“intersexual niche partitioning hypothesis”), or may be result from differential selection among the sexes on maximizing reproductive output (“sexual selection hypothesis”). The latter may lead to higher energy demands in females driven by fecundity selection, while males invest in mate searching. We tested predictions of the two hypotheses underlying intersexual trophic niche partitioning in a natural population of spiders. Zodarion jozefienae spiders specialize on Messor barbarus ants that are polymorphic in body size and hence comprise potential trophic niches for the spider, making this system well-suited to study intersexual trophic niche partitioning.

Methodology/Principal Findings

Comparative analysis of trophic morphology (the chelicerae) and body size of males, females and juveniles demonstrated highly female biased SSD (Sexual Size Dimorphism) in body size, body weight, and in the size of chelicerae, the latter arising from sex-specific growth patterns in trophic morphology. In the field, female spiders actively selected ant sub-castes that were larger than the average prey size, and larger than ants captured by juveniles and males. Female fecundity was highly positively correlated with female body mass, which reflects foraging success during the adult stage. Females in laboratory experiments preferred the large ant sub-castes and displayed higher capture efficiency. In contrast, males occupied a different trophic niche and showed reduced foraging effort and reduced prey capture and feeding efficiency compared with females and juveniles.

Conclusions/Significance

Our data indicate that female-biased dimorphism in trophic morphology and body size correlate with sex-specific reproductive strategies. We propose that intersexual trophic niche partitioning is shaped primarily by fecundity selection in females, and results from sex-differences in the route to successful reproduction where females are selected to maximize energy intake and fecundity, while males switch from foraging to invest in mating effort.     

Sexual dimorphism and sexual segregation in foraging strategies of northern giant petrels, Macronectes halli, during incubation

Giant petrels ( Macronectes spp.) are the most sexually dimorphic of all seabirds. We used satellite-tracking and mass change during incubation to investigate the influence of sexual size dimorphism, in terms of the intersexual food competition hypothesis, on foraging and fasting strategies of northern giant petrels at South Georgia. Females foraged at sea whereas males foraged mainly on the South Georgia coast, scavenging on seal and penguin carcasses. Foraging effort (flight speed, distance covered, duration of foraging trips) was greater for females than for males. In contrast, foraging efficiency (proportionate daily mass gain while foraging) was significantly greater for males than for females. Females were significantly closer to the desertion mass threshold than males and could not compensate for the mass loss during the incubation fast while foraging, suggesting greater incubation costs for females than for males. Both sexes regulated the duration and food intake of foraging trips depending on the depletion of the body reserves. In males the total mass gain was best explained by mass at departure and body size. We suggest that sexual segregation of foraging strategies arose from size-related dominance at carcasses, promoting sexual size dimorphism. Our results indicate that sex-specific differences in fasting endurance, contest competition over food and flight metabolic rates are key elements in maintenance of sexual size dimorphism, segregating foraging strategies and presumably reducing competition between sexes.     

Foraging behaviour of non-breeding Western Sandpipers Calidris mauri as a function of sex, habitat and flocking

Individuals within a population may vary considerably in the way they exploit available food resources. If the sexes differ in the size of their feeding apparatus, there can be differences in foraging behaviour and habitat use, hence one sex may be more susceptible to competition. We examined relationships between sexual dimorphism in bill size and foraging behaviour, and habitat and microhabitat use of non-breeding Western Sandpipers Calidris mauri at Bahía Santa María, northwestern Mexico. Western Sandpipers are sexually dimorphic, with females about 15% longer-billed than males. Males used a higher proportion of pecks, had a lower probing–pecking rate, walked at higher rates, foraged at sites with lower water content, and had greater variation in foraging technique than females. Moreover, males decreased their proportion of pecks and foraged at a higher rate than females when they changed from feeding alone to feeding in flocks, suggesting a greater safety advantage or susceptibility to conspecific interference when flock feeding. We compared behaviour and microhabitat usage in three habitats: brackish flats, mangroves, and cattail marshes. Sex-related differences in foraging behaviour and microhabitat use were consistent among habitats. Birds in brackish flats and mangroves used a higher proportion of pecks, foraged at lower rates and walked at higher rates, and foraged at deeper sites, with a lower proportion of water cover, than those in cattail marshes. Sex-related differences in foraging behaviour and microhabitat should reduce the level of competition between sexes, and may account for sex differences in Western Sandpiper distribution observed between habitats in Bahía Santa María.     

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.     

Ecomorphological relationships in a group of insectivorous birds of temperate forests in winter

We examined the relationships between morphology and foraging behaviour in a group of insectivorous birds wintering in temperate mixed forests in northern Iberia. Using principal components analysis we reduced 11 biometric variables to three major morphological components and 20 foraging categories to four major ecological factors. The relative length of the tarsometatarsus and bill morphology were the most important morphological variables predicting foraging ecology. Birds exploiting distal parts of trees and foliage were generally smaller and had relatively longer tarsometatarsi than those foraging on trunks. Foraging on the ground and branches of medium diameter was associated with bill thickness. Ecomorphological patterns were discernible at the level of substrate use and foraging methods, but bear no relation to selection of tree species or foraging height. Morphology correctly predicted niche breadth and interspecific overlap. In Parus spp. interspecific differences in bill shape could explain 63% of the interspecific segregation according to substrate use (R²= 0.63, p <0.01).     

同域分布3种啄木鸟冬季取食的生态位差异

为了掌握黑啄木鸟、三趾啄木鸟和大斑啄木鸟的冬季取食行为特征,特别是三者之间取食生态位的差异,于2016年1月5-13日,在黑龙江省凉水国家级自然保护区以样线法结合样方法对3种啄木鸟的取食生境和取食行为进行了系统调查,收集了15个生境和行为特征变量数据。共布设45条样线,484个对照样方,收集312组啄木鸟取食数据,其中黑啄木鸟73组,三趾啄木鸟97组,大斑啄木鸟142组。多变量回归树和多分类逻辑斯谛分析结果显示,3种啄木鸟在所调查的15项特征上存在显著分异。采用基于利用-可利用方法的Bailey''s方法和双因子方差分析,分别对3种啄木鸟的生境选择和行为特征进行分析,结果显示：黑啄木鸟和三趾啄木鸟偏好在郁闭度较高的原始云、冷杉林中取食,而大斑啄木鸟则随机地在各种林型、生境中取食。黑啄木鸟、三趾啄木鸟多在树干取食,黑啄木鸟更常在倒木上取食,而大斑啄木鸟则多在树冠层取食。黑啄木鸟基本只在主干上凿洞,其他两种特别是大斑啄木鸟则可以在侧枝上取食。与黑啄木鸟和大斑啄木鸟凿洞取食昆虫不同,三趾啄木鸟多通过扒去树皮获得食物。黑啄木鸟的取食树基本为死树,单树取食时间最长,大斑啄木鸟多在活树上取食,单树取食时间最短,经常更换取食树,而三趾啄木鸟的取食树则死活参半,单树取食时间也较长。黑啄木鸟的冬季取食行为节律表现为双峰形,日出后和日落前各有一个活动高峰,其他两种则于白天持续取食。3种啄木鸟取食生境和行为生态位的差异,使它们能够更有效地利用有限的食物资源,共存于同一森林。     

Seasonal dimorphism in the horny bills of sparrows

Bill size is often viewed as a species‐specific adaptation for feeding, but it sometimes varies between sexes, suggesting that sexual selection or intersexual competition may also be important. Hypotheses to explain sexual dimorphism in avian bill size include divergence in feeding niche or thermoregulatory demands, intrasexual selection based on increased competition among males, or female preference. Birds also show seasonal changes in bill size due to shifts in the balance between growth rate and wear, which may be due to diet or endogenous rhythms in growth. Insight into the function of dimorphism can be gained using the novel approach of digital x‐ray imaging of museum skins to examine the degree to which the skeletal core or the rhamphotheca contribute to overall dimorphism. The rhamphotheca is ever‐growing and ever‐wearing, varying in size throughout life; whereas the skeletal core shows determinant growth. Because tidal marsh sparrows are more dimorphic in bill size than related taxa, we selected two marsh taxa to investigate dimorphism and seasonality in the size of the overall bill, the skeletal core, and the rhamphotheca. Bill size varied by sex and season, with males having larger bills than females, and bill size increasing from nonbreeding to breeding season more in males. Skeletal bill size varied with season, but not sex. The rhamphotheca varied primarily with sex; males had a larger rhamphotheca (corrected for skeletal bill size), which showed a greater seasonal increase than females. The rhamphotheca, rather than the skeletal bill, was responsible for sexual dimorphism in overall bill size, which was particularly well developed in the breeding season. The size of the rhamphotheca may be a condition‐based character that is shaped by sexual selection. These results are consistent with the evidence that bill size is influenced by sexual selection as well as trophic ecology.     

Ecological causes for the evolution of sexual dimorphism: a review of the evidence

Can sexual dimorphism evolve because of ecological differences between the sexes? Although several examples of this phenomenon are well known from studies on birds, the idea has often been dismissed as lacking general applicability. This dismissal does not stem from contradictory data so much as from the difficulties inherent in testing the hypothesis, and its apparent lack of parsimony, in comparison to the alternative explanation of sexual selection. The only unequivocal evidence for the evolution of sexual dimorphism through intersexual niche partitioning would be disproportionate dimorphism in trophic structures (e.g., mouthparts). This criterion offers a minimum estimate of the importance of ecological causes for dimorphism, because it may fail to identify most cases. A review of published literature reveals examples of sexually dimorphic trophic structures in most animal phyla. Many of these examples seem to be attributable to sexual selection, but others reflect adaptations for niche divergence between the sexes. For example, dwarf non-feeding males without functional mouthparts have evolved independently in many taxa. In other cases, males and females differ in trophic structures apparently because of differences in diets. Such divergence may often reflect specific nutritional requirements for reproduction in females, or extreme (sexually selected?) differences between males and females in habitats or body sizes. Ecological competition between the sexes may be responsible for intersexual niche divergence in some cases, but the independent evolution of foraging specializations by each sex may be of more general importance. If ecological causation for dimorphism can be demonstrated in so many cases, despite the inadequacies of the available criteria, the degree of sexual size dimorphism in many other animal species may well also have been influenced by ecological factors. Hence, it may be premature to dismiss this hypothesis, despite the difficulty of testing it.     

Sex promotes spatial and dietary segregation in a migratory shorebird during the non-breeding season

Several expressions of sexual segregation have been described in animals, especially in those exhibiting conspicuous dimorphism. Outside the breeding season, segregation has been mostly attributed to size or age-mediated dominance or to trophic niche divergence. Regardless of the recognized implications for population dynamics, the ecological causes and consequences of sexual segregation are still poorly understood. We investigate the foraging habits of a shorebird showing reversed sexual dimorphism, the black-tailed godwit Limosa limosa, during the winter season, and found extensive segregation between sexes in spatial distribution, microhabitat use and dietary composition. Males and females exhibited high site-fidelity but differed in their distributions at estuary-scale. Male godwits (shorter-billed) foraged more frequently in exposed mudflats than in patches with higher water levels, and consumed more bivalves and gastropods and fewer polychaetes than females. Females tended to be more frequently involved and to win more aggressive interactions than males. However, the number of aggressions recorded was low, suggesting that sexual dominance plays a lesser role in segregation, although its importance cannot be ruled out. Dimorphism in the feeding apparatus has been used to explain sex differences in foraging ecology and behaviour of many avian species, but few studies confirmed that morphologic characteristics drive individual differences within each sex. We found a relationship between resource use and bill size when pooling data from males and females. However, this relationship did not hold for either sex separately, suggesting that differences in foraging habits of godwits are primarily a function of sex, rather than bill size. Hence, the exact mechanisms through which this segregation operates are still unknown. The recorded differences in spatial distribution and resource use might expose male and female to distinct threats, thus affecting population dynamics through differential mortality. Therefore, population models and effective conservation strategies should increasingly take sex-specific requirements into consideration.     

Effects of body size and social context on the arboreal activities of lowland gorillas in the Central African Republic

The objectives of this 27 month study were to document the positional behaviors used by lowland gorillas (Gorilla gorilla gorilla) in the Central African Republic and to compare the effects of body size dimorphism on the use of arboreal substrates. During this study, despite their great size, all gorillas used trees regularly. Predictions concerning the relationship of body size to arboreal behavior were generally upheld. Small branch and suspensory activities were rare for silverbacks. Females used smaller and multiple substrates and suspensory postures more frequently than males. Although females foraged in the periphery of trees, males stayed close to the cores and rarely used terminal branches. In addition to body size, this study found that party size, social rank, and tree structure all influence an animal's substrate choice and subsequent positional activities. Lone males typically remained in the cores of trees where substrates are large. Group males may have been forced to use all parts of trees because others were present. Lone males used small crown trees which provided easy access to terminal branch foods. Males and females foraging together used larger trees (containing more feeding sites) than single sex groups. Female positional behavior may have been affected by the presence of males. When apart from males, females used the cores of trees and larger substrates more than when foraging with males. As habitat and social context both influence substrate use, they should be considered essential components of body-sized based interpretations of the behavior of fossil or extant species. © 1995 Wiley-Liss, Inc.     

Reversed sexual dimorphism in tawny owls, Strix aluco, correlates with duty division in breeding effort

Even though most bird species with a raptorial feeding habit express varying extents of reversed sexual dimorphism (RSD: females bigger than males), the evolutionary basis for its maintenance, as well as its possible secondary consequences for the ecological adaptations of the different sexes, is debated. We studied pairs of tawny owls, Strix aluco (females 20% heavier than males), throughout the year by telemetry to test whether any inter-sexual differences in movement patterns, resource partitioning and breeding effort correlated with RSD. Females were larger than males in all body size measures and were 16% heavier than would be expected from the difference in wing length alone. In accordance with predictions from flight economics, males moved longer distances per time unit than females, in particular during the post-fledging season, when they also fed chicks more often than the females. Males had larger home ranges than females during the post-fledging period, whereas the sexes had home ranges of equal size during the non-breeding season. Until 10 days after fledging, females foraged much closer to the offspring than males, apparently balancing their distance to offspring between the needs of offspring guarding and foraging. In males, the parent–offspring distance only increased with decreasing brood condition. The sexes did not differ in habitat use or feeding habits, rendering no indications of food niche partitioning. The study provides further evidence that selection for males to be light and energetically efficient foragers is the main evolutionary force behind RSD in raptorial birds, even when the prey base is confined by territoriality.     

Foraging Behaviour of Subordinate Great Tits (Parus major). Can Morphology Reduce the Cost of Subordination?

This paper studies the magnitude of the behavioural shift, from forage standing to forage hanging, of subordinate great tits ( Parus major ) in two different social contexts: feeding solitarily vs. feeding with a dominant conspecific. The aim is to test the hypothesis that differences in morphological design provide subordinates with varying abilities to reduce the presumed costs of subordination. We find that different subordinate individuals change the foraging behaviour, occupying a different niche when an intra-specific competitor is present. Morphology linked to sexual dimorphism, specifically body mass, is the factor responsible for the different magnitudes of change. Lighter subordinates can remain longer than heavier ones at the feeding patch without interrupting their foraging. Thereby, the former reduce the costs of being subordinate more than the latter. Among subordinates, females are lighter than males; they also spend more time feeding in the presence of a dominant conspecific than males do. No differences are found between age categories. We find no relationship between tarsus length and individual ecological plasticity. Our results support the idea that the ecological plasticity due to morphological differences is a mechanism that allows subordinate individuals to overcome costs associated with subordination.     

灰椋鸟身体大小和内脏器官形态的两性差异

鸟类性二态现象广泛存在,比如身体大小、羽色等,性二态很可能是自然选择和性选择共同作用的结果.为了探索和更好地了解雀形目鸟类身体大小性二态的进化,在2019年繁殖季节早期研究了灰椋鸟(Sturnus cineraceus)野外种群身体大小和内脏器官形态的两性差异.结果表明,除嘴宽外,其他身体特征参数均雄性显著大于雌性,表...     

Sexual dimorphism in bill morphology and feeding ecology in Cory's shearwater (Calonectris diomedea)

The bill is a sexually dimorphic structure in many bird species and implicated in numerous functions. Sexual differences may arise from sexual selection or ecological divergence. Here, we examined differences in bill size and shape between males and females and explored to what extent these relate to feeding ecology of each sex in Cory's shearwater (Calonectris diomedea). We applied linear measurements and geometric morphometric methods to examine sexual differences in bill size and shape. We investigated feeding ecology by tracking foraging movements during the breeding period and by analysing stable isotope signatures in blood during the breeding period and in feathers grown during the non-breeding period. Bill traits were all sexually dimorphic, both in absolute and relative terms, and scaled hypermetrically with body mass in several characters in males. However, males and females did not differ in their feeding areas or isotopic signatures and no significant correlation was observed between these traits and bill dimorphism. Therefore, we discard the foraging-niche divergence hypothesis, and suggest that sexual dimorphism in bill size in this species is more likely driven by sexual selection related to antagonistic interactions.