The reproductive behavior of Anthidium manicatum (Hymenoptera: Megachilidae) and the significance of size for territorial males

Summary Males of the wool-carder bee, Anthidium manicatum, patrol clumps of garden plants. Females of this species visit these plants for pollen, nectar, and pubescence; they also mate there. Females are polyandrous, with intervals between copulations as short as 35 s. Patrolling males defend their territories (0.1–1.3 m²) against other males and against other species of flower-visiting insects. Honey bees may be rendered unable to fly by the attacks of A. manicatum.Territory owners perform exploratory flights to other males' territories, changing territories often (median ownership 4–7 days; maximum 30 days) and flying up to 450 m to establish new territories. Territorial usurpations are nearly always by larger males.Female visitation rate is significantly correlated with number of flowers on a territory. The head size of territory-owner males shows significant correlation with territorial quality (measured by number of flowers, not area) and thus with number of female visits and copulatory opportunities. Some males fail to maintain territories and instead attempt to forage and copulate in other males' territories while the owners are otherwise occupied. Nonowner males are significantly smaller than owners, forage less often and from fewer flowers, and achieve significantly fewer copulations than owners. Females, however, do not reject smaller, nonowner males at a higher rate than they do larger, owner males; their choice for male size appears to be indirect, based instead on choice of food resource.The interval between a copulation and the male's next attempt with a different female is not shorter than that involving the same female. Males do not escort just-mated females about their teritories, as observed in Anthidium maculosum. Territorial behavior in this species most likely evolved through intrasexual competition for reproductive success which led to sexual dimorphism. The defense of a resourcebased territory is the mechanism used by a male to maximize his reproductive potential.     

Nectar dehydration by male carpenter bees as preparation for mating flights

Summary Females of all social and many solitary bees dehydrate nectar before storing it or adding it to larval provisions. Nectar dehydration by males has rarely been documented. We report on the neotropical facultatively social carpenter bee Xylocopa nigrocincta, in which the nest constant males are fed nectar by their female nestmates. Males dehydrate the nectar at the nest entrance before leaving the nest for mating territories. We show that males thereby minimize their water load, resulting in an improvement of their energy budget during hovering flights in their territories. Males can prolong the duration of territorial flights if they cany highly concentrated nectar. We assume that nectar dehydration as a pre-mating behavior is not correlated with the social organization in Xylocopa species. However, the behavior is particularly weil-developed in X. nigrocincta, where during the mating period males remain integrated in the nest society and are fed by their mothers and sisters.     

The role of distance,familiarity, and time of day in Carolina wrens responses to conspecific songs

Summary We tested two predictions of the ranging hypothesis (Morton 1982) which suggests that: (1) For distance estimation of singing intruders from them, birds compare degraded songs they hear to undegraded songs in their memory. (2) The strength of response of birds to songs depends on how well they can range the songs as coming from inside or outside the territory.In accordance with first prediction we found that, Carolina wrens discriminate significantly between songs played from outside to those played inside their territories only if they are familliar with the song type played (Fig. 1). In accordance with the second prediction, on the average, the birds in our study responded stronger to unfamiliar than to familiar songs when these were played from outside the territory, but stronger to familiar than to unfamiliar songs when played inside it (Fig. 2). We concluded that this was because they could range the familiar songs with greater certainty.The variance in the data was, in part, due to changes in the birds' responses during the day. We report for the first time, an increase in the strength of territorial defense of birds later in the morning, accompanied by a decrease in singing in response to song playback. We suggest that these changes reflect an increase in energy gain and worse acoustic conditions later in the day.     

When do helpers help? Food availability and helping in the moorhen,Gallinula chloropus

Summary The importance of food availability as a constraint upon cooperative behaviour was examined by providing supplementary food on some breeding territories of a population of moorhens (Gallinula chloropus). First brood juveniles on territories provided with food were more likely to feed their younger second brood siblings. Supplementary feeding improved the physical condition of juveniles above that of unfed conspecifics, and birds in better condition were more likely to be helpers. Helpers on fed territories also provided more food to siblings than did their counterparts on unfed territories, enabling parent birds to reduce their work rates. Results are discussed in relation to the benefits which may accrue to helpers and the energetic costs of cooperation which may act as proximate constraints upon potential helpers.     

Contrasting foraging tactics by northern gannets (<Emphasis Type="Italic">Sula bassana</Emphasis>) breeding in different oceanographic domains with different prey fields

In order to forage and to provision offspring effectively, seabirds negotiate a complex of behavioural, energetic, environmental and social constraints. In first tests of GPS loggers with seabirds in North America, we investigated the foraging tactics of free-ranging northern gannets (Sula bassana) at a large and a medium-sized colony that differed in oceanography, coastal position and prey fields. Gannets at Low Arctic colony (Funk Island) 50 km off the northeast coast of Newfoundland, Canada provisioned chicks almost entirely with small forage fish (capelin Mallotus villosus, 89%), while at boreal colony (Bonaventure Island) 3 km from shore in the Gulf of St. Lawrence, Quebec, Canada, large pelagic fish dominated parental prey loads (Atlantic mackerel Scomber scombrus 50%, Atlantic herring Clupea harengus 33%). Mean foraging range and the total distance travelled per foraging trip were significantly greater at the larger inshore colony (Bonaventure) than at the smaller offshore colony (Funk Island; 138 and 452 km vs. 64 and 196 km, respectively). Gannets from Funk Island consistently travelled inshore to forage on reproductive capelin shoals near the coast, whereas foraging flights of birds from Bonaventure were much more variable in direction and destination. Birds from the Low Arctic colony foraged in colder sea surface water than did birds from the boreal colony, and dive characteristics differed between colonies, which is concordent with the difference in prey base. Differences between the colonies reflect oceanographic and colony-size influences on prey fields that shape individual foraging tactics and in turn generate higher level colony-specific foraging “strategies”.     

Territoriality among male red-winged blackbirds

Summary We examined male site fidelity and territorial movements in a population of individuallyidentifiable red-winged blackbirds (Agelaius phoeniceus) over an 8-year period. Based on general theory and knowledge of the ecology of this population, we predicted that males would be site conservative, and that they would voluntarily change territories only when they could expect to improve significantly their reproductive success. Because males are absent from their territories for only short periods, and probably have accurate comparative information only about nearby territories on which to base their decisions, we predicted that most moves would be over short distances. About 70% of males that bred for more than 1 year retained their original territories between breeding years. Most of the 30% of males that changed territories moved less than 200 m, often to adjacent territories. As predicted, males moving less than 200 m tended to have better reproductive success after moving than before, whereas long distance movers did not improve their success after moving. Territorial male redwings appear to monitor breeding activity on nearby territories and move when significant potential benefits are perceived and opportunities exist. Long-distance moves may be involuntary ones. The amount of information possessed about present and alternative sites, and the time and opportunity required to collect the information, are probably major constraints influencing site conservatism in this and other territorial species.     

Begging and short-term need in cowbird nestlings: how different are brood parasites?

Because brood parasitic nestlings are usually unrelated to their nestmates and to the provisioning adult, they are free from indirect costs of begging. Consequently, they are predicted to beg more intensely than host nestlings, and some models predict they will beg at an invariantly high level, regardless of short-term need. Previous work has shown intense begging by parasitic cowbirds, but short-term need was not controlled. In this study, we manipulated short-term need and measured begging intensity in two species pairs of host and parasitic nestlings: shiny cowbirds (Molothrus bonariensis) parasitizing larger hosts than themselves (rufous-bellied thrushes, Turdus rufiventris), and brown-headed cowbirds (M. ater) parasitizing smaller hosts than themselves (yellow warblers, Dendroica petechia). All four species increased their begging intensity with short-term need, though the change was much less pronounced between food-deprived and control treatments than between control and hand-feeding treatments. Shiny cowbirds begged more intensely than rufous-bellied thrushes following each treatment. In contrast, brown-headed cowbirds did not beg significantly more intensely than yellow warblers under any of the treatments. Intense begging by both species of parasites was as effective as host begging in stimulating the adults to make provisioning visits, but shiny cowbirds were less successful at acquiring food from adult thrushes. A wide array of factors may underlie the apparent differences in the begging behavior and success of brown-headed and shiny cowbirds, including relative size, experience, and local risk of nest predation. Our experiments clearly demonstrate, however, that these two species of parasitic cowbirds adjust their begging intensity based on short-term need.Communicated by H. Kokko     

Echolocation,development, and vocal communication in the lesser bulldog bat,Noctilio albiventris

Summary Foraging and echolocation behavior and its ontogeny in the lesser bulldog bat, Noctilio albiventris, were studied in Panama under field and captive conditions. The vocalizations utilized for echolocation and communication were monitored. Adult N. albiventris captured insect prey from the water surface employing various combinations of CF/FM (constant frequency and frequency modulated) signals. The proportions of CF/FM and the repetition rate were a function of the bat's activity. Most adults exhibited post-sunset and pre-dawn foraging activity, although several telemetered lactating females foraged for only the half hour after dusk, spending the rest of the night with their babies in the roost. When the juveniles began to leave the roost at the age of two months, they appeared to accompany their mothers on initial flights.Captive infant Noctilio developed slowly, and did not fly until about 5–6 weeks postnatally. They continued to nurse for almost 3 months, even though they were capable of eating solid food at about 6 weeks. Previous to weaning, mothers fed their infants with masticated food from their cheekpouches.At birth, Noctilio emit a combination of long FM isolation calls and shorter CF/FM pulses. Mothers nurse only their own babies which they appear to recognize by a vocal signature contained in the infants' isolation calls. The individual isolation calls, as well as the mother's communication sounds, appear to be variations of an FM sinusoidal wave. The periodicity and amplitude change, and different portions including harmonics are added or deleted. The short CF/FM signals of the infant evolve into the adult orientation type signals as the CF component increases in frequency and the repetition rate increases. These sounds appear to serve a dual function in communication and echolocation. Mother-young pairs were observed to call antiphonally, utilizing CF/short FM signals in retrieval situations. This duetting was also observed in bats flying over the Chagras River after the time the juveniles began to fly, and may function to maintain vocal contact during initial foraging flights.Deceased     

Correlation between nectar supply and aggression in territorial honeyeaters: causation or coincidence?

Summary A fundamental prediction of food-based economic models of territoriality is that animals will not defend territories if food is so abundant that defense will not improve access to food. Several studies of nectar-feeding birds support this prediction, with territoriality being rare or absent in years when nectar was particularly abundant. However, these results could potentially be an artefact of changes in bird density with nectar availability, and in at least some cases the correlations between territory defense and nectar availability could be purely coincidental. This paper reports the first experimental test of whether cessation of territory defense in nectar-feeding birds results from a direct response to abundance of nectar. New holland honeyeaters Phylidonyris novaehollandiae and white-cheeked honeyeaters P. nigra show pronounced changes in their levels of territorial aggression over the 7–8 months that they breed. These changes are predictable from economic considerations in that the birds are least aggressive in the months when nectar is extremely abundant. I tested whether the birds were responding to changes in nectar availability by providing sugar-water feeders at neutral locations that were easily accessible to territory holders, but far enough away from territories that intrusion rates were unaffected. I tested for responses at two time scales feeders were put out for 48-h periods in 1987, and were left out continuously from January to October 1988. The only effect was that territory holders visited feeders instead of flowers when floral nectar was scarce. They continued to defend their territories aggressively at those times, showed seasonal changes in aggressiveness similar to birds on a site without feeders, and did not shift their territories toward feeders. I conclude that the observed changes in aggressiveness are not responses to changes in nectar availability, and suggest alternative explanations.     

Stopover behaviour and departure decision of northern wheatears, Oenanthe oenanthe, facing different onward non-stop flight distances

On the small North Sea island Helgoland (54°11' N, 07°55' E) we studied the stopover ecology of two subspecies of northern wheatear, Oenanthe oenanthe, during spring migration. Birds heading for Scandinavia (O. o. oenanthe) face only short flights across an ecological barrier (50-500 km) whereas those originating from Greenland and Iceland (O. o. leucorhoa) have to cover between 1,000 and 2,500 km in the impending flight. Colour-ringed individuals showed that 90% of Scandinavian birds left on the day of ringing while 40% (males) and 30% (females) of Greenland/Icelandic birds stayed at least 1 night. The birds who remained were thus mostly O. o. leucorhoa. They often established desirable feeding territories on the beach and had a high rate of body mass increase (1.7 g/day). However, subspecies did not differ in habitat choice and in foraging effort, but O. o. leucorhoa had a higher success rate in pecking. Departure decisions were analysed by comparing (a) conditions on the day of ringing between departing and staying birds and (b) for birds staying between the day of departure and the preceding day. The factors that were probably important in the decision to depart differed between subspecies. In O. o. leucorhoa, few birds departed with bad or deteriorating weather conditions (tailwind component, cloud cover), whereas departures of O. o. oenanthe seemed to be little affected by those factors. A few O. o. oenanthe stayed early in the spring migration season and/or had low fat reserves. Interference during foraging seemed to play a role because both subspecies tended to leave when the densities of northern wheatears were high. Other factors related to refuelling conditions (food supply, foraging effort, predation risk) failed to show differences between staying and departing individuals. In summary, almost all Scandinavian birds departed quickly and irrespective of refuelling and weather conditions, whereas many (but not all) Greenland/Icelandic birds seemed to prepare for a long-distance flight and carefully adjusted departure to weather conditions. The observed differences in stopover behaviour and departure decisions in the two subspecies of northern wheatear indicate that the distance to the next stopover site or to the goal area has to be considered when applying optimal migration models.     

Orientation and navigation in bats: known unknowns or unknown unknowns?

Bats have been extensively studied with regard to their ability to orient, navigate and hunt prey by means of echolocation, but almost nothing is known about how they orient and navigate in situations such as migration and homing outside the range of their echolocation system. As volant animals, bats face many of the same problems and challenges as birds. Migrating bats must relocate summer and winter home ranges over distances as far as 2,000 km. Foraging bats must be able to relocate their home roost if they range beyond a familiar area, and indeed circumstantial evidence suggests that these animals can home from more than 600 km. However, an extensive research program on homing and navigation in bats halted in the early 1970s. The field of bird navigation has advanced greatly since that time and many of the mechanisms that birds are known to use for navigation were not known or widely accepted at this time. In this paper I discuss what is known about orientation and navigation in bats and use bird navigation as a model for future research in bat navigation. Technology is advancing such that previous difficulties in studying orientation in bats in the field can be overcome and so that the mechanisms of navigation in this highly mobile animal can finally be elucidated.     

Spatio-temporal use of the urban habitat by feral pigeons (Columba livia )

Feral pigeons are descendants of wild rock pigeons that have adapted to the urban habitat. They have partially conserved the foraging behaviour of their wild ancestors (flights to agricultural areas) but have also developed new habits. Previous studies on the foraging strategies of feral pigeons have given various results, e.g. maximum distances reached by the pigeons (measured in a straight line from the resting places) differed between 0.3–0.5 km and 18–25 km. This study focuses on the spatio-temporal activity of feral pigeons in the urban habitat. We equipped 80 free-living feral pigeons from Basel, Switzerland with GPS receivers. We found three different foraging strategies for pigeons in Basel: (1) in the streets, squares and parks near the home loft, (2) in agricultural areas surrounding the city, (3) on docks and railway lines in harbours. The maximum distance reached by a pigeon was 5.29 km. More than 32% of the pigeons remained within 0.3 km of the home lofts and only 7.5% flew distances of more than 2 km. Females covered significantly longer distances than males, preferring to fly to more abundant and predictable food sources. Temporal activity patterns showed to be influenced by sex, breeding state and season. In contrast to wild rock pigeons and to feral pigeons in other cities, pigeons in Basel showed a clear bimodal activity pattern for breeding birds only. The differences between our results and those of other studies seem to be partly method-dependent, as the GPS-technique allows to record the pigeons’ localisations continuously in contrast to other methods. Other differences might be due to different kinds of food supply in the various cities. Our study shows that feral pigeons have individual foraging strategies and are flexible enough to adapt to different urban environments.Electronic supplementary material Supplementary material is available for this article at     

Territorial fidelity of willow warbler (Phylloscopus trochilus) males and success in competition over territories

Summary Territorial fidelity and competition between males was studied in a population of individually marked willow warblers (Phylloscopus trochilus). After spending the winter in Africa males arrive in the study area in late April or the beginning of May. They can be separated into two categories, old colour-banded males that return for a new breeding attempt, and birds that are new to the study area. These new birds could either be birds that had bred the previous year returning to a different site, or more likely 1-year-old birds returning for their first breeding attempt. Old males show high site fidelity and are successful in retaining their territories from the preceding year. The few old males that switched territories between years seemed to do so in response to reduced breeding success in the preceding year. However, in 13 out of 14 cases when a new resident was already established on the previous resident's territory, the old male was successful in evicting the new resident. In a removal experiment both old and new birds were removed from their territories and then released when a replacement male had been established. All old males successfully managed to reclaim their original territories after escalated contests. None of the new males were successful and no fights were observed.     

Individual variation in mate care by alpha males in crested tit winter flocks

Summary Winter flocks of crested tits Parus cristatus, typically consisting of two adults and one or two non-kin 1st-year birds, were observed to split up into subflocks in a way related to ambient temperature. On warm days, when they were foraging in subflocks without 1 st-year birds, alpha males always occupied the most preferred upper foraging sites, as expected from their top dominance rank. On cold days, when foraging in flocks with 1st-year birds, 8 out of 13 alpha males shifted to lower (less preferred) positions below their alpha mates while allowing the latter to forage at the best sites. As enhanced access to preferred microsites on days with high energy stress is believed to increase overwinter survival probability, this shifting behaviour of alpha males can be considered as a form of mate care. Out of 13 alpha males, however, 5 did not shift and always occupied the best foraging sites irrespective of flock composition. As (i) these non-shifting males were in poorer physical condition than shifting males, (ii) they scanned significantly less for predators than either females or shifting males when foraging in the uppermost tree parts, and (iii) four out of five non-shifting males were replaced by immigrants in early spring, absence of mate care during winter may be caused by constraints due to condition. High-quality-territory owners in poor condition at the end of autumn were most vulnerable to replacement by immigrants. Therefore, as four out of five replacements affected high-quality territories, selective intrusion by immigrants is suggested. Correspondence to: L. Lens     

Body morphologies shape territorial dominance in the satyrine butterfly <Emphasis Type="Italic">Lethe diana</Emphasis>

Male butterflies compete over mating territories via aerial interactions. It has often been suggested that flight performance, and thus flight-related traits, would play an important role in butterflies’ contest resolution; however, most studies have not demonstrated a clear relationship between these traits and territory ownership in butterflies. Males of the satyrine butterfly Lethe diana compete over territories via linear chases, a behavior that is uncommon among butterflies: rather, contests of most butterfly species consist of circling flights. In the present study, some morphological and physiological traits of L. diana were compared between territory owners and intruders to search for traits correlated with resource holding power (RHP). Territory owners had higher flight-muscle ratio (FMR) and were heavier than intruders, indicating that flight performance plays a key role in their contest resolution. Contest duration was not affected by the owners’ FMR or body mass, and thus I obtained no evidence supporting the hypothesis that contestants assess their relative RHP before making a decision to retreat. I speculate that the fact that these traits had an effect on territorial status in L. diana was a result of the characteristics of the contest behavior of this butterfly. During their contests, males L. diana fly faster in their linear flights than males of most other butterflies, which engage in slower circling flights. Therefore, males of L. diana probably need higher flight performance capabilities in order to win their contests.     

Migratory fuelling in blackcaps (Sylvia atricapilla) under perceived risk of predation

It has been argued that the body mass levels achieved by birds are determined by the trade-off between risks of starvation and predation. Birds have also been found to reduce body mass in response to an increased predation risk. During migration, the need of extra fuel for flights is obvious and crucial. In this study, migratory blackcaps (Sylvia atricapilla) were subject to an experimental stopover situation where the predation risk was manipulated by exposure to a stuffed predator. Blackcaps that perceived an imminent risk of predation increased their food intake and fuel deposition rate during the first period of stopover compared with a control group. The pattern of night activity indicates that birds that were exposed to the predator also chose to leave earlier than birds in the control group. Since there was no cover present at the stopover site, birds might have perceived the risk of predation as higher regardless of whether they were foraging or not. Under such circumstances it has been predicted that birds should increase their foraging activity. The findings in this study clearly indicate that birds are able to adjust their stopover behaviour to perceived predation risk. Received: 8 January 1997 / Accepted after revision: 11 April 1997     

The adaptive significance of male subadult plumage in purple martins: plumage dyeing experiments

Summary I tested two hypotheses for the adaptive significance of subadult plumage in male purple martins (Progne subis) : the female mimicry and subordinance signaling hypotheses. Subadult males were at a competitive disadvantage in obtaining territories, as they arrived later in the spring than adult males. Contrary to the predictions of both hypotheses, adult male territory owners were not less aggressive toward subadult male than adult male intruders. The subadult plumage was not effective in mimicking females, as adult male owners were significantly more aggressive toward subadult male than female intruders. Summer adaptation hypotheses predict that young males in subadult plumage are more successful in acquiring territories and mates than they would be with an adult plumage. I tested this prediction by dyeing the plumage of floater subadult males to mimic the appearance of adult males. In 13/17 paired experiments, dyed subadults obtained territories before control subadults. There was no difference in the time it took dyed and control males to attract a mate after they obtained a territory. These results suggest that the subadult plumage is not an advantage to young males in competing with adult males for breeding resources. In late winter, subadult males were growing mostly femalelike feathers on their underside, suggesting that the subadult plumage is not the result of a molt constraint. The subadult plumage could enhance survival of yearlings in winter roosts if it improves access to good roost sites or reduces the risk of predation.     

Size and spacing of capercaillie leks in relation to social behavior and habitat

Summary Spring territories of 17 adult capercaillie (Tetrao urogallus L.) cocks were 10–79 ha in extent; they varied inversely in size with the relative proportion of mature forest within them. The number of resident cocks at leks increased with the amount of mature forest within a 1 km radius of each lek center. Leks in two areas (n=46) were regularly spaced with mean distances to nearest neighbors of 1.98 and 2.07 km, corresponding to the territorial space occupied by adult cocks of adjoining leks. In one area with intensive logging, interlek distance increased with decreasing amounts of mature forest between them. Spring home ranges of 18 adult females averaged 51.3 ha±8.2 SE. The spatial relationships did not fit recent models of interlek spacing based on female spacing behavior. Instead, the results suggested that spacing of leks may be related to the territorial requirements of males.     

Blue eggs do not reduce nest predation in the song thrush,Turdus philomelos

Summary Many passerine birds with open cup-shaped nests lay blue or blue-green eggs. In thrushes, blue eggs may be cryptic and provide camouflage by imitating spots of light on green leaves. Alternatively, egg coloration may be selectively neutral because nest predators detect nests and not eggs, or it may be maladaptive because organisms are not always well adapted to their present environment. I evaluated these hypotheses by studying predation on artificial song thrush (Turdus philomelos) nests with quail eggs, painted either white, blue, or spotted (cryptic to a human eye). Corvids were the major nest predators. For concealed as well as exposed nests, I found no differences in the predation rates of nests with white, blue, or spotted eggs. Predators apparently detected the nests, and not the eggs, first. In a second experiment, I placed egg groups without nests in trees to study the effect of color per se. The predation rate of the spotted egg groups was significantly lower than that of the white and blue egg groups, for concealed as well as exposed egg groups. These results suggest that blue eggs in the song thrush are not cryptic but may be selectively neutral or even maladaptive with regard to nest predation.     

Territorial behavior of the black and yellow rockfish and gopher rockfish (Scorpaenidae,Sebastes)

Tagged black-and-yellow rockfish Sebastes chrysomelas and gopher rockfish S. carnatus were observed in situ for 1 yr at 3 study sites off California, USA in order to determine the nature of their home ranges and territories. Three basic patterns of space use were identified: (1) a shelter hole located within a larger area of activity (in which the fish presumably fed, since its size increased with size of fish and decreased with prey density); (2) discontinuous home ranges, in which the shelter holes and feeding areas were separte; and (3) dispersed home ranges with little or no exclusive area. Fish in the first category were regarded as territorial, since their home ranges were defended and relatively exclusive. Because individuals did not patrol borders, however, territories often overlapped at their peripheries. In addition, some territory holders appeared to tolerate the sheltering of other fish within their territories, as long as these intruders left the territory during periods of activity. Some fish with discontinuous home ranges (commuters) may have been territorial, but others may have been forced to occupy suboptimal feeding areas. Fish in the third category (floaters) were regarded as nonterritorial; their activities were apparently limited to areas where they could escape the aggression of territory holders. The varying patterns appeared to develop through defense and accommodation during the turnover of individuals, with each fish securing shelter and feeding areas as these became available.