Territorial Males Research Articles

Thermoregulatory Requirements Shape Mating Opportunities of Male Proboscis Bats

The spatiotemporal distribution of females is a major factor affecting animal social systems. Predation risk and the distribution of feeding resources often determine where females are found, but abiotic factors (e.g. temperature) can also shape the distribution of females and therefore variation in social organization and mating systems. Given the predicted future changes in climatic variation, it is vital to understand how animal mating systems and the sexual selection process may be altered by temperature. In bats, female distribution is tightly linked to roosting ecology and particularly to the microclimatic conditions at the roost. Proboscis bats (Rhynchonycteris naso) form cohesive and stable multi-male-multi-female groups and inhabit exposed day roosts (e.g. tree trunks, vines, buildings). Strong selection to remain inconspicuous to visually oriented predators in the exposed day roosts has been suggested to promote a rather rare male mating strategy termed site-specific dominance where males defend females directly but are successful in doing so only in their own territory. The choice of open-roost structures can result in the bats roosting under direct sunlight, making individuals susceptible to overheating. Here we investigate whether regular relocations of R. naso social groups among male territories are a mechanism of behavioral thermoregulation. Our results suggest that in general R. naso choose the warmest suitable roost sites within a roost, possibly to minimize the energetic costs of thermoregulation. However, on days with high midday temperatures at the primary roost site, bats commonly relocate to alternative, cooler sites within their roosts. These thermoregulatory relocations entail that a social group regularly switches among the territories of several males. Thus, the need for behavioral thermoregulation determines the spatial distribution of females and shapes the mating opportunities of males during the day. This is supported by our result that territorial males defending primary roost sites are reproductively more successful than territorial males of alternate roost sites. In line with other studies, our findings suggest that the increase in ambient temperatures associated with climate change has the potential to affect the intensity of sexual selection in bat species and may have far-reaching behavioral, demographic and evolutionary consequences for their populations.

Territorial patterns of Dickerson’s collared lizard, Crotaphytus dickersonae

High quality males generally obtain better territories, resulting in enhanced survival rates and reproduction. In our study we examined which phenotypic traits play a role in obtaining the best territories in males of the highly territorial and sexually dimorphic lizard species Crotaphytus dickersonae . For each lizard, we measured snout-vent length, body mass, head width, hind limb length, and tail length as phenotypic traits that might promote success in competition over territories. In addition, we examined whether the bright blue coloration of the males is related to territory quality. We assessed territory quality by determining the size, the number of overlapping female home ranges, the exclusivity of each territory, and the number of refuges and basking sites. Males with longer tails and wider heads had the best territories with more females and less overlapping male territories. Bluer males tended to have less overlapping male territories and more overlapping female home ranges, suggesting that the blue coloration is related to male quality. Since tail length and head width influence fighting ability in lizards, males with longer tails and wider heads probably obtain the best territories through male-male competition.

Dominance hierarchy establishment in the invasive round goby, Neogobius melanostomus

Organisms living at high densities may be forced to engage in conflict for access to resources such as food or shelter. When these resources are limited, the outcome of interactions may have important fitness implications. We investigated the behavioural interactions of the invasive Round Goby (Neogobius melanostomus) in a shelter-limited environment. Round Goby are benthic fish that utilize rocky shelters for predator avoidance and as reproductive sites in which territorial males defend clutches of eggs. Previous work on this and other species has shown that larger individuals have greater resource holding potential in dyadic interactions. In order to understand the outcome of agonistic interactions in more complex social environments, we observed groups of three goby of the same sex which varied in relative size in an aquarium in which individuals had the opportunity to compete for access to shelters. We predicted that larger goby would behave aggressively towards smaller goby, and outcompete smaller goby for access to shelters. Because males defend shelters while breeding, we also predicted that male goby would compete more aggressively than females over dominance status. We found that larger goby in groups were socially dominant to smaller goby, regardless of sex. Additionally, we found that the largest goby in each group was involved in more aggressive interactions than the second or third largest goby in each group. We found no effect of relative size or sex on aggressive interaction or the emergent dominance relationships. Our findings highlight that aspects of the social environment may limit the opportunity for individuals to establish dominance or establish ownership of resources.

Male competition drives song divergence along an ecological gradient in an avian ring species.

Sexual selection operates via female choice and male competition, which can act independently, in concert, or in opposition. Female choice is typically considered the stronger selective force, but how these two processes interact to shape phenotypic divergence is poorly understood. I tested the hypothesis that variation in sexual selection in different habitats drives song divergence in the greenish warbler ring species. I evaluated the strength, direction, and targets of female choice and male competition in three populations spanning 2400km of latitude. Average song length increased with latitude, concomitant with a decline in population density. Within populations, males sang longer songs when females were fertile and shorter songs during territory establishment. Females consistently preferred males with longer songs and larger song repertoires. By contrast, playback experiments showed that males used short songs in territory defense. Songs were shortest at high densities, and in the highest density population only, song traits preferred by females correlated with male territoriality. Stronger male competition at high population densities likely constrains maximum song length, whereas weaker competition at low densities allows expression of female choice for long songs. Interactions between male competition and ecology may be a crucial but oft-overlooked component of phenotypic divergence and speciation.

Influence of military training on breeding ecology of Bachman's sparrow

ABSTRACTAnthropogenic disturbance may cause birds to flush and relocate, abandon breeding sites, experience increased nest failure, or fledge fewer young. Ground‐based military activities are of particular concern for ground‐nesting birds because of the increased risk of nest destruction and trampling of vegetation. We investigated how different intensities of disturbance from ground‐based military training affected reproductive ecology of Bachman's sparrow (Peucaea aestivalis) from 2014–2016, on Fort Bragg Military Installation, North Carolina, USA. We designated 2 training intensity regimes and monitored sparrows at 6 observation areas, 3 in high‐intensity training areas (i.e., foot traffic every 1–3 days) and 3 in low‐intensity training (i.e., foot traffic <1 per month) areas. We compared seasonal productivity metrics and daily nest survival between training intensities. Additionally, we compared male sparrow relative abundance and micro‐habitat use between high‐ and low‐intensity training areas. We monitored 106 male territories and located 110 nests opportunistically and by tracking telemetered female sparrows. We used fixed‐radius point counts to estimate relative abundance in each observation area and measured vegetation composition and structure at a subset of 10 locations in each male territory. Seasonal productivity metrics, daily nest survival, relative abundance, and vegetation composition and structure at male locations did not differ between areas with high and low military training intensity. In 2015, 1 sparrow nest was trampled by military personnel, but ≥1 nestling force‐fledged and the nest was considered successful. Bachman's sparrow nesting ecology was not affected by intensity of ground‐based military training activity, likely because activity was sufficiently dispersed across the landscape, even in high‐intensity training areas. We recommend ground‐based training attempts to minimize ground vegetation destruction by dispersing large groups and traversing variable routes when in forested uplands. © 2018 The Wildlife Society.

Song performance is a condition‐dependent dynamic trait honestly indicating the quality of paternal care in the bull‐headed shrike

The good parent hypothesis in sexual selection predicts that if females can increase their fitness by mating with males who provide high‐quality parental care, then female preferences for male phenotypes honestly indicating the quality of paternal care will evolve. In order to test this hypothesis, we investigated correlations between male song, the timing of pair formation of males, male feeding rate, and reproductive success, in the bull‐headed shrike Lanius bucephalus (an altricial oscine passerine with biparental care). Analysis revealed that although the timing of pair formation was not correlated with most song characteristics (e.g. repertoire size), male morphological traits (e.g. tail length), or male territory size, it was negatively correlated with male singing tempo (i.e. the number of notes uttered per second). Those males that sang at higher speeds were in better body condition (i.e. body mass standardized by tarsus length3 was higher), fed their chicks frequently during the nestling period, and raised heavy chicks. These results show that in the bull‐headed shrike male singing tempo is a condition‐dependent dynamic trait honestly indicating the quality of paternal care, and strongly supports the good parent hypothesis.

When to change your tune? Unpaired and paired male house wrens respond differently to anthropogenic noise

In response to anthropogenic noise, many bird species adjust their song frequency, presumably to optimize song transmission and overcome noise masking. But the costs of song adjustments may outweigh the benefits during different stages of breeding, depending on the locations of potential receivers. Selection might favor unpaired males to alter their songs because they sing to attract females that may be widely dispersed, whereas paired males might not if mates and neighbors are primary receivers of their song. We hypothesized male house wrens (Troglodytes aedon) respond differently to noise depending on their pairing status. To test our hypothesis we synthesized pink noise, which mimics anthropogenic noise, and played it at three intensities in territories of paired and unpaired focal males. We recorded their songs and analyzed whether song structure varied with pairing status and noise treatment. To validate our study design, we tested whether noise playback affected measurement of spectral song traits and changed noise levels within territories of focal males. Consistent with our predictions, unpaired males sang differently than paired males, giving longer songs at higher rates. Contrary to predictions, paired males changed their songs by increasing peak frequency during high intensity noise playback, whereas unpaired males did not. If adjusting song frequency in noise is beneficial for long-distance communication we would have expected unpaired males to change their songs in response to noise. By adjusting song frequency, paired males reduce masking and produce a song that is easier to hear. However, if females prefer low frequency song, then unpaired males may be constrained by female preference. Alternatively, if noise adjustments are learned and vary with experience or quality, unpaired males in our study population may be younger, less experienced, or lower quality males.

Surviving in the city: higher apparent survival for urban birds but worse condition on noisy territories

AbstractAnthropogenic landscapes and soundscapes impose strong selective pressures on a number of species, which can manifest in changes in vocalizations, foraging strategies, predator vigilance, and reproductive success. However, few studies have examined survival rates, a major component of fitness, across urban landscapes and soundscapes. White‐crowned sparrows (Zonotrichia leucophrys) persist in both urban and rural landscapes and change their behavior in response to the soundscape. We color‐banded adult white‐crowned sparrows and collected noise levels on their territories in the urban San Francisco Bay Area and adjacent rural Point Reyes National Seashore in California. We collected mark‐encounter data on territorial males from 2014 to 2017. Using the Program MARK, we tested the effects of habitat (urban/rural) and territory noise level on annual survival rates and body condition. We predicted that survival and body condition would be lower in urban habitats and decrease with increasing background noise level on territories. We found that survival estimates vary according to year, and males in urban landscapes have higher survival. Noise levels best predict body condition, such that soundscape negatively correlates with male body condition. Taken together, the urban landscape and soundscape shape the survival and health of birds in and near cities.

Black British Women Filmmakers in the Digital Era: New Production Strategies and Re-Presentations of Black Womanhood

Abstract The story of Black women in British mainstream cinema is certainly one of invisibility and misrepresentations, and Black women filmmakers have historically been placed at the margins of British film history. Up until the mid-1980s, there were no Black female directors in Britain. Pioneers like Maureen Blackwood, Martina Attille and Ngozi Onwurah have actively challenged stereotypical representations of Black womanhood, whilst asserting their presence in Black British cinema, often viewed as a male territory. In the 2010s, it seems that the British film industry remains mostly white and masculine. But the new millennium has brought a digital revolution that has enabled a new generation of Black women filmmakers to work within alternative circuits of production and distribution. New strategies of production have emerged through the use of online crowdfunding, social media and video-sharing websites. These shifts have opened new opportunities for Black women filmmakers who were until then often excluded from traditional means of exhibition and distribution. I will examine these strategies through the work of Moyin Saka, Jaha Browne and Cecile Emeke, whose films have primarily contributed to the re-presentation of Black womanhood in popular culture.

Ritualised Dung Kicking by White Rhino Males Amplifies Olfactory Signals but Reduces Odour Duration.

Many mammals enhance their olfactory signals visually by depositing them in conspicuous locations such as well-travelled paths. It is also possible to enhance the odour itself through behaviours aimed at modifying odour emission rates. White rhinos defecate in communal middens. While defecating, territorial males kick sharply with their back feet which disperses their dung. Despite being a ubiquitous trait of territorial male white rhinos, the reason behind this behaviour is unclear. We hypothesised that the purpose of dung kicking was for olfactory signal amplification (OSA) in terms of an increased emission of volatile compounds (i.e. increased signal strength). Using dung collected from non-territorial adult males (because it is not possible to collect whole dung from territorial males), we show that the dispersal of male white rhino dung causes OSA by increasing the emission of hydrocarbon acids. The dung odour of territorial and non-territorial males differs only quantitatively, hence it is likely that the same emission patterns occur for territorial male dung odours following dung dispersal. The volatile compound indicating age of intact dung was toluene, but for dispersed dung it was acetophenone (similar to territorial male dung). Despite the benefits of OSA, dung dispersal carried a cost of decreased odour longevity. Thus, signal detectability is temporally reduced. However, territorial males likely counteract this by defecating in middens both before and during peak visitation times by other individuals (15:00-23:00). As a result, we suggest that dung kicking by territorial males amplifies signal strength, such that their dung odours are the most prominent and easily detectable by individuals visiting the middens. This would then better signal territorial ownership to both potential rivals and potential mates.

Territorial behaviour in southern red-backed and Ozark zigzag salamanders: effects of sex, species and ownership

Abstract When both males and females are territorial, spatial and aggressive behaviour can vary between the sexes. We examined differences in exploratory and competitive behaviour between males and females and residents and intruders of two species, the Ozark zigzag (P. angusticlavius) and southern red-backed (P. serratus) salamanders. For exploratory behaviour, males and females differed in distance traveled based on species, but females of both species were significantly more cautious than males about leaving territories. In competition trials, male and female territory residents (owners) did not differ in defensive behaviour, but female intruders were more aggressive than male intruders. Overall, P. angusticlavius showed more aggressive behaviour, explored more, and were less cautious about leaving territories than P. serratus. Female intruders of both species fight harder, potentially to gain territories, and are more cautious about leaving territories, which may indicate that the cost of not having a territory is greater for females.

Song function and territoriality in male and female White-throated Dippers Cinclus cinclus

ABSTRACTCapsule: Male White-throated Dippers Cinclus cinclus are more likely and quicker to respond to the playback of song than females, but both sexes are more likely to respond before the onset of breeding than after.Aims: Territoriality and the function of song in female birds have rarely been studied outside of the tropics or Australasia. We investigated territoriality and song function in males and females of a Northern temperate species, the White-throated Dipper.Methods: We conducted playback trials on established pairs and compared the responses of males and females according to the sex of the simulated intruder and the timing of playback relative to the onset of breeding. A response was classified as movement towards the speaker, singing or both.Results: Males were significantly more likely and quicker to respond to playback than females, but neither sex responded differently to the playback of male and female song. Both sexes were more likely to respond to playback before breeding had begun than after.Conclusions: Our results suggest that both males and females are territorial but that males take the dominant role in defence. Female song appears to elicit a similar response to male song and may play a role in territoriality or mate defence.

Brilliant-thighed poison frogs do not use acoustic identity information to treat territorial neighbours as dear enemies

Some territorial animals recognize familiar neighbours and are less aggressive to established neighbours than they are to strangers. This form of social recognition produces a ‘dear enemy’ effect, which may allow animals to reduce the costs of territory defence. The dear enemy effect is thought to reflect either the decreased threat posed by neighbours relative to strangers (the relative threat hypothesis) or the decreased need for escalation with increasing familiarity between neighbours (the familiarity hypothesis). We tested for a vocally mediated dear enemy effect in male brilliant-thighed poison frogs, Allobates femoralis. In this species, the familiarity hypothesis predicts a dear enemy effect, because males defend long-term stable territories and should be familiar with the calls of their neighbours. In contrast, the relative threat hypothesis does not predict a dear enemy effect, because neighbours and strangers both represent competitors for mates and likely pose equivalent threats. Acoustic analyses showed that males produce individually distinctive advertisement calls. Two playback experiments were conducted to determine whether territorial males respond less aggressively to neighbours' calls than to strangers' calls based on this acoustic identity information. In the first experiment, males pursued acoustically simulated neighbours and strangers to similar distances when calls were played both from the direction of the neighbour's territory and from a direction with no neighbours. In the second experiment, males responded aggressively to neighbours' and strangers' calls at similar threshold amplitudes and pursued these calls to similar distances. Hence, two different playback experiments failed to find evidence of a vocally mediated dear enemy effect. Our results support the hypothesis that territorial animals respond to the relative threat posed by neighbours and strangers regardless of their level of familiarity with neighbours.

Female preference for males that have exclusively marked or invaded territories depends on male presence and its identity in the subterranean rodent Ctenomys talarum

AbstractTerritorial scent‐marking provides chemical records of male competitive interactions that are available to females, who gain valuable information to assess and identify best quality partners. In this context, the solitary subterranean rodent tuco‐tuco (Ctenomys talarum) offers excellent possibilities to evaluate the effects of male exclusive scent‐marking of territories on female assessment. For evaluation, we used wild caught individuals of C. talarum, manipulated their scent marks within the territories in captive conditions and staged preference tests where females were able to choose between exclusive and invaded territories. The evaluation was performed in two scenarios considering the identity of the intruder scent mark: territories invaded by a strange male and territories invaded by a neighbour male. Females investigated the chemical cues deposited on the substrate of the exclusively marked territory more frequently. Next, females displayed equal interest to scent samples of both males presented in a Y‐maze. Finally, when females could gain access to both individually isolated males and their scent‐marked territories, they spent more time within invaded territories despite they visited them with the same frequency. Moreover, females tried to get in contact by scratching the mesh of the owner of the invaded territory more frequently. We found that females of C. talarum evaluate the homogeneity (exclusiveness) of scent marks within a male territory and then show preferences in relation to the identity of the intruder's scent –whether strange or neighbour.

The role of middens in white rhino olfactory communication

White rhinos, Ceratotherium simum, use dung odours to transmit information about their sex, age, territorial status (males) and oestrous state. Moreover, as white rhinos defecate in communal middens (i.e. dung heaps, or latrines) it has been suggested that these middens may act as information centres. However, it is uncertain which individuals primarily transmit information via middens, or for whom this information is intended. Using video-recording camera traps, we investigated the behaviour of white rhinos at middens. We hypothesized that territorial adult males would visit, defecate and sniff other dung more than other adults. In line with this, we found that they visited and defecated more than other individuals. Moreover, territorial males and potential male challengers were the main individuals to investigate dung piles. These olfactory investigations focused primarily on territorial male and adult female dung (male–male and female–male communication). Although investigating less often, investigation by adult females and subordinate males was also focused on territorial male and female dung, suggesting male–female and female–female communication. In addition to olfactory signals, there was a spatial aspect to midden use, where territorial males defecated only in the centre of a midden, while other individuals defecated primarily around the periphery. Yet, subordinate males also tended to defecate in the centre, suggesting an indication of residency. Lastly, territorial males defecated more frequently than any other adult, and were able to do so by regulating their dung output (i.e. producing smaller volumes per deposit). Our results indicate that middens act as information centres, where the primary function seems to be for territorial males to transmit and obtain information. However, nonterritorial males may also assess female reproductive state, while females may be assessing the quality of all males, and the number of other females using a midden. Ultimately, our results highlight the importance of middens in white rhino communication.

Queuing, takeovers, and becoming a fat cat: Long‐term data reveal two distinct male spatial tactics at different life‐history stages in Namibian cheetahs

AbstractIn mammals, male spatial tactics and sociality can be predicted from the size, degree of overlap and ease of defense of female individual or group home ranges (HRs). An exception is apparently the cheetah (Acinonyx jubatus) for which studies from East Africa describe a socio‐spatial organization where resident males defend small territories, which cover a portion of large ranges of solitary females, and non‐territorial males (floaters) roam over vast areas whilst queuing for access to territories. Most studies from southern Africa did not report the existence of territorial males and floaters, but a system with both males and females roaming over vast areas with overlapping HRs. Here, we derive and test predictions from previously described spatial tactics in felids by studying the movements, behavior, and/or physical characteristics of 164 radio‐collared Namibian cheetahs on commercial farmland from 2002 to 2014. The results demonstrate the existence of male territory holders and floaters and a, by mammalian standards, unique sociality in that commonly groups of males, sometimes solitary males defended small areas partially overlapping with large ranges of solitary females. When a solitary male or a group of males switched between both tactics, floating usually preceded territory holding, suggesting that both spatial phases are equivalent to distinct life‐history stages. Switching from roaming as a floater to holding a territory was also associated with an increase in body mass index (BMI) and a change in the observed behavior of animals captured in traps when approached by humans. Both BMI and this behavior are therefore reliable, quick biomarkers of an individual's space use tactic and life‐history stage. We elaborate the implications of this socio‐spatial organization for models of ecological movements and on conflict mitigation measures such as translocations or the planning of future protected areas. We suggest that such implications also apply to other species where one sex exhibits two space use tactics and two sets of range sizes.

Aging male loons make a terminal investment in territory defense

Animals that senesce experience a decline in residual reproductive value (RRV), such that old individuals can expect reduced breeding success compared to young ones. According to life history theory, animals with low RRV, which thus have less to lose, should shift resources away from self-maintenance and towards reproduction, an expectation called terminal investment. In a population of common loons whose survival and territorial behavior were measured throughout life, males 14 years and older exhibited clear senescence, as they showed lower survival, reduced body mass, and far greater susceptibility to territorial eviction than younger males. While older males invested no more effort than young males in feeding or protecting their chicks, they increased territorial yodeling by 35%, showed more aggression towards territorial intruders, and, following eviction from original territories, resettled with great frequency on vacant, unproductive territories nearby. Our findings thus provide support for terminal investment in territorial behavior. Hyper-aggressive behavior by old, declining males might explain the unusual occurrence of lethal combat for territories in this species. Life history theory holds that animals that decline with age should invest less energy in staying alive and more in reproduction when they become old, because they have less to lose in doing so. Evidence for such terminal investment, however, is scanty. We measured rates of survival and territorial behavior in a known-age population of common loons and found that males, but not females, suffered a decrease in survival rate and ability to hold a territory after age 14. At the same time, territorial males older than 14 behaved more aggressively and were more apt to give a territorial yodel call towards territorial intruders. Hence, our study appears a rare example of terminal investment in aggressive behavior, whereby old, declining males cling desperately to their territories in hopes of producing a few more offspring before they die.

Soft songs in male ortolan buntings are used in an aggressive context but are not an aggressive signal

AbstractMany bird species produce low‐amplitude acoustic signals that have been poorly studied in comparison with loud, broadcast songs used for mate attraction and repelling rivals. In some birds, these soft signals were found to be emitted in an antagonistic context and were the most reliable predictor of a subsequent physical attack. The function of this signal is poorly understood, and several hypotheses have been proposed to explain the mechanisms and possible functions of such low‐amplitude signals. The subject of this study is the ortolan bunting, a small passerine species that produces soft songs during territorial defence. In this study, we aim to study whether the soft songs of the ortolan bunting are a signal of increased aggressiveness by testing if they meet the context, prediction and response criteria of aggressive signals. We simulated stranger male intrusion into a focal male territory with three different playback experiments. We found no significant differences in the male responses to the taxidermic model regardless of whether they were or were not producing soft songs in response. The males responded more strongly to loud songs than to soft songs during the simulated intrusions, and the males did not treat soft songs as a predictor of conflict escalation. Although soft songs clearly appeared during territorial encounters and were not present during spontaneous singing before the intrusions, our results did not support the hypothesis that soft songs indicate aggressive character. We suggest that soft songs in the ortolan bunting are intentionally used by birds to modify their intentions or target‐specific individuals within a close range.

A nocturnal rail with a simple territorial call eavesdrops on interactions between rivals.

The behaviour of most animals has evolved in a communication network environment, in which signals produced by senders are perceived by many intended and unintended receivers. In this study, we tested whether the corncrake (Crex crex), a nocturnal rail species with innate (non-learned) calls, is able to eavesdrop on the interactions of conspecific males and how this eavesdropping affects subsequent responses by the eavesdropper to territorial intrusion. In the first step, simulated aggressive or neutral interactions between male dyads were presented to a focal male. In the second step, the calls of winning, losing or neutral males from the first step were played within the territory of the focal male. We measured behavioural and vocal responses of focal males. We found that corncrakes eavesdropped on signal exchange between rivals. Males often began responding to distant aggressive interactions during the eavesdropping phase, and they responded strongly during the intrusion phase of the experiments. The response was significantly weaker to playback of males from neutral interactions than to those involved in aggressive interactions, and we found no differences between the responses to Winners and Losers entering a focal male territory.

Effects of Supplemental Food on the Behaviour and Paternity Status of Male Indigo Buntings (Passerina CyaNea)

Breeding is energetically costly and, if energy-limited, birds may alter their time budgets, spending less time engaged in some activities and more time in others. Investigators who have provided breeding birds with supplemental food have noted changes in time budgets, but the extent and types of changes have been found to vary among species. Our objective was to determine how food supplementation might influence the time budgets and paternity status of male Indigo Buntings (Passerina cyanea). We studied Indigo Buntings from 8 May to 15 August 2009 in Madison County, Kentucky. Territories of male Indigo Buntings (n=30) were randomly assigned as either food supplemented (n=8) or controls (n=22). During time-budget observations, all behaviours were noted. To determine paternity status, blood samples were collected from adults and nestlings at 16 nests. Males with feeders in their territories spent significantly less time foraging and more time vocalising (chip notes) than males without feeders. Time spent singing and mate guarding by treatment and control males did not differ. Differences in the proportion of extra-pair young did not differ between nests in territories with and without supplemental food. Our results suggest that, when provided with supplemental food, male Indigo Buntings spend less time foraging and more time engaged in other activities. However, neither the presence of supplemental food nor differences in the behaviour of males in food-supplemented versus non-supplemented territories affected rates of extra-pair paternity.