Food availability,intrusion pressure and territory size: an experimental study of Anna's hummingbirds (Calypte anna)

Theoretical considerations implicate food availability and intrusion pressure as important determinants of territory size, but empirical studies have led to contradictory conclusions about cause-and-effect relationships among these three variables. To investigate this problem, we provided patches of electronically controlled artificial flowers, which were defended by male Calypte anna. Food availability was experimentally manipulated, and intrusion rate and territory size were calculated from behavioral observations of the territory owner. Changes in both food availability and intrusion rate were found to be significantly correlated with changes in territory size under certain conditions. Intrusion rate, which was influenced by food availability, was negatively associated with territory size so long as food availability was high. This association persisted even after possible effects of food availability were controlled statistically. Food availability was negatively correlated with territory size only when intrusion rates were high and after owners had been defending territories for 3 days. As food availability and intrusion rate increased, owners increasingly restricted their defense to the patch itself; partial regressions revealed a significant association for intrusion rate but not food availability. When intrusion rate was low and food availability varied from low to high levels, no relationship was observed between food availability and territory size, apparently because of opposing influences of food abundance on territory size. Correspondence to: P.W. Ewald     

Effect of territory quality on intrusion rate in nonbreeding hummingbirds

Summary Experimental reductions of productivity on feeding territories of Anna's hummingbirds (Calypte anna) decreased intrusion rate. This trend contrasts with the absence of strong correlations between intrusion rate and productivity in previous experimental studies. Consideration of sample sizes, productivity rates, and time-dependent influences on intrusion rate (e. g., day-to-day changes in weather) indicates that this difference resulted from the more natural spatial distribution of artificial flowers used in this study. Offprint requests to: P.W. Ewald     

Territory quality and territorial behavior in two sympatric species of hummingbirds

Summary Changes in territorial behavior of blackchinned hummingbirds (Archilochus alexandri) in response to experimental changes in territory quality were investigated using artificial feeders and simultaneous, pair-wise observations of owners. Some of the responses of A. alexandri were similar to those documented by a previous study of the Anna's hummingbird (Calypte anna): A. alexandri defending rich territories spent more time on their territories, encountered a greater percentage of intruders, and chased more intruders than did A. alexandri defending poor territories. In contrast to C. anna, A. alexandri supplemented chases with energetically inexpensive threat vocalizations more extensively when territory quality was increased. This difference may be related to A. alexandri's more tenuous control of rich territories. When both species were observed on very poor territories, A. alexandri chased a greater percentage of intruders, consumed a greater proportion of available food, and obtained a greater net energy gain from their territories. When observed simultaneously on territories with ad lib food, both species defended highly exclusive territories but A. alexandri suffered higher intrusion pressure and spent more time and energy in defense. These interspecific differences in territorial behavior may facilitate sympatric coexistence of the two species through aggressive partitioning of flower patches according to productivity: the greater payoff received by C. anna on rich territories and A. alexandri on poor territories should favor a corresponding monopolization of rich areas by C. anna and poor areas by A. alexandri.     

Traplining by purple-throated carib hummingbirds: behavioral responses to competition and nectar availability

We examined the effects of nectar availability and competition on foraging preferences and revisit intervals of traplining female purple-throated caribs hummingbirds (Eulampis jugularis) to Heliconia patches shared by two individuals or visited solely by one individual. Birds at both shared and solitary patches preferred multiflowered to single-flowered inflorescences, but the magnitude of this preference depended on food availability and competition. During a year of low flower availability, females visited multiflowered inflorescences more frequently than single-flowered inflorescences only when nectar availability was experimentally enhanced; similarly, females at shared patches exhibited a significant preference for multiflowered inflorescences only after experimental increases in nectar availability. Experimental manipulations of nectar availability also had different effects on revisit intervals of birds at shared vs solitary patches. Birds at shared patches responded to patch-wide increases in nectar rewards by increasing the duration of their visit intervals, whereas birds at solitary patches did not. In contrast, birds at solitary patches responded to abrupt losses of nectar at flowers (simulating competition) by decreasing the duration of their visit intervals, whereas a bird at a shared patch did not alter its return interval. The contrasting results between shared vs solitary patches suggest that future studies of traplining behavior should incorporate levels of competition into their design.     

Thermoregulatory cost affects territorial behavior in hummingbirds: a model and its application

A common assumption in behavioral ecology is that the valuation of a resource by consumers depends on the energetic value of the resource itself. Nevertheless, the value of a resource may be relative to the condition of the organism, which is in turn related to the abiotic conditions such as ambient temperature. We developed a theoretical model—incorporating these untested assumptions—to predict a functional relationship between territorial aggression and ambient temperature for individuals sensitive to daily variations in energy availability. We evaluated our theoretical predictions against a field experiment carried out with the hummingbird Sephanoides sephaniodes. The model predicted a quadratic relation between aggression intensity and ambient temperature. Field data were better explained by a quadratic equation than a linear function, suggesting the existence of lower and upper thresholds of temperature which determine the intensity of territorial defense. Ambient temperature affects energy expenditure for thermoregulation, and therefore, it fixes the benefit level that must be produced by the territory to pay the costs of its defense. Our findings strongly suggest that abiotic conditions can change an animal evaluation of the yield of a resource and in turn influence the behavioral strategy which it adopts.     

Effects of territory quality, food availability and sibling competition on the fledging success of oystercatchers (Haematopus ostralegus)

We investigated the fledging probability of oystercatcher, Haematopus ostralegus, chicks as a function of hatching order, brood size, territory quality and food availability. Sibling dominance was related to the hatching order in both low- (’leapfrogs’) and high-quality (’residents’) territories. Differences in hatchling mass might have aided the establishment of a dominance hierarchy, since breeders produced small late eggs and hatchlings. These mass differences were most pronounced in leapfrogs, and in large broods in years with lower food availability (’poor’ years). Late hatchlings fledged less often and with lower body masses compared to early hatchlings in all situations. Leapfrogs produced smaller broods and hatched their broods more asynchronously in poor years than leapfrogs breeding in years with more available food (’good’ years) and residents breeding in both poor and good years. Large brood sizes resulted in lower survival of hatchlings in poor years. These results favour the ’brood reduction’ hypothesis. However, contrary to the expectations of this hypothesis, hatching order also affected fledging success in residents. Moreover, large brood size resulted in higher survival of hatchlings in good years, particularly in residents. Thus, although large broods experienced losses due to sibling competition in some years, they nevertheless consistently produced more fledglings per brood in all years, both as leapfrogs and residents. We believe this effect is due to parental quality correlating with initial brood size. Most leapfrogs, at best, fledged one chick successfully each year, losing chicks due to starvation. Nevertheless, leapfrog broods were reduced in size after hatching significantly less quickly than resident broods. These results suggest that breeders lay and hatch insurance eggs to compensate for unpredictable losses due to the high predation rates on both nests (ca 50%) and chicks (ca 90%), in accordance with the ’nest failure’ hypothesis. Received: 14 February 2000 / Revised: 27 September 2000 / Accepted: 10 June 2000     

Temperature effects on behavior and survival of marine invertebrates exposed to variations in hydrostatic pressure

Investigation into the pressure resistance of 8 intertidal species belonging to the genera, Uca, Sesurma, Talorchestia and Littorina from the Northern Gulf of Mexico and Panama sheds light on the relationships between temperature and hydrostatic-pressure resistance. Generally, increasing temperature increases the pressure required to elicit reversible reactions such as increased activity and tetany, or paralysis, wheroas increasing temperature generally evokes the irreversible response of death (LD₅₀) at a decreasing pressure. Tropical stenotherms tend to be more sensitive to hydrostatic pressure than eurythermal-temperate species at the same or similar temperatures.     

Determinants of social dominance and inheritance of agonistic behavior in an island population of silvereyes,Zosterops lateralis

Summary We examined components of agonistic behavior and dominance in parents and offspring of silvereyes on Heron Island, Great Barrier Reef, to discover their determinants. The data were collected over four years on 1,235 juveniles and their parents. Of four behavioral characters used to compare juveniles and their parents, bill clattering was seen significantly more among young from female parents that exhibited bill clattering. The total absence of submissive behavior and the total absence of aggressive behavior were shown more often than not by young from dominant male parents and subordinate female parents, respectively. Young fledging early in the breeding season tended to be more dominant than those fledging late in the season. The number of brood-mates or the local nest density did not affect the dominance of young. Dominance status appears to be little affected by the performance of parents.     

Mate competition in the velvet swimming crab Necora puber: effects of perceived resource value on male agonistic behaviour

Male velvet swimming crabs [Necora puber (L.)] were observed to engage in long and vigorous agonistic interactions to gain possession of a sexually receptive female. The role of agonistic behaviour in male mate competition in this species was examined by investigating the effect of the perceived presence of sexually receptive females on interactions between males. Exposing male crabs to water conditioned by sexually receptive females resulted in prolonged interactions, with a greater incidence of potentially injurious behaviour than interactions between control crabs, exposed only to sea water. Maleconditioned water also resulted in more offensive behaviour, but these interactions were of shorter duration and not significantly different from sea water controls. Agonistic superiority was strongly correlated with contestant relative size when males were exposed to male-conditioned water or sea water, but not when exposed to female-conditioned water. Overall, the results conformed with the general predictions of game theoretic models in relation to the influence of resource value on agonistic behaviour. Crabs were more persistent and probably incurred greater fitness costs in the perceived presence of a sexually receptive female, when interactions may have been resolved on the basis of factors more closely related to actual fighting ability than the relative body sizes of contestants.     

Age,coloration and dominance in nonbreeding hummingbirds: A test of the asymmetry hypothesis

Summary During the nonbreeding season, adult Anna and black-chinned hummingbirds (Calypte anna and Archilochus alexandri) have lower defense costs and more exclusive territories than juveniles. Adult C. anna are victorious over juveniles in aggressive encounters, and tend to monopolize the most temporally predictable resources.Juveniles are more successful than adults at stealing food from territories (the primary alternative to territoriality), presumably because juveniles are less brightly colored. Juveniles have lighter wing disc loading than adults, and consequently should have lower rates of energy expenditure during flight. Reduced flight expenditures may be more important for juveniles because their foraging strategy requires large amounts of flight time. These results support the contention of the asymmetry hypothesis that dominance can result from a contested resource being more valuable to one contestant than to the other.Among juveniles, defence costs are also negatively correlated with age and coloration; amount of conspicucus coloration is negatively correlated with the number of bill striations, an inverse measure of age.     

Density dependence and habitat quality modulate the intensity of display territory defence in an exploded lekking species

We evaluated the effect of conspecific abundance and habitat quality of leks on the territorial behaviour of males in an exploded lekking species, the Little Bustard (Tetrax tetrax). The hypothesis that males more intensely defend territories with higher conspecific abundance and better habitat quality was evaluated experimentally analysing the agonistic response of experimental males to male decoys placed on their displaying areas. Decoy experiments showed that the intensity of display territory defence by little bustard males is density dependent. The time experimental males took to return to their display sites after decoy placement decreased with abundance of both males and females. The strength of their final response was positively associated to local male and female abundance in the vicinity of their display sites. Habitat quality also influenced males’ display territory defence since the intensity of male response increased with the degree of natural vegetation cover. Habitat quality was particularly relevant in explaining variation of experimental males’ snort call rate, which decreased with the degree in plough cover and increased with the number of fields in the lekking area. Snort call rate decreased with the level of male aggregation and was lowest in males exhibiting the strongest aggressive response to decoys. These results add new evidence for the density dependence of species’ breeding territorial behaviour, supporting density-dependent models of lek formation and reinforcing the role of resources defence in exploded lek mating systems.     

Relative importance of male and territory quality in pairing success of male rock ptarmigan (Lagopus mutus)

We studied pairing success in male rock ptarmigan (Lagopus mutus) in northern Alaska to learn whether males obtaining more females possessed phenotypic traits that influenced female choice directly, whether these traits permitted males to obtain territories favored by females, or whether both processes occurred. The number of females per male varied from zero to three. Several male and territory traits were significantly correlated with number of females per male. We used multiple regression to obtain a single measure of male quality and a single measure of territory quality. These measures of male and territory quality correlated with each other and with male pairing success. We used path analysis to separate direct effects of male quality on pairing success from indirect effects due to high-quality males obtaining high-quality territories. Both direct and indirect pathways had significant effects on pairing success, and direct and indirect effects of male traits on pairing success were about equal. This study illustrates an analytical approach for estimating the relative importance of direct and indirect causal relationships in natural systems. Received: 13 January 1998 / Accepted after revision: 7 November 1998     

Yolk androgens in great tit eggs are related to male attractiveness,breeding density and territory quality

Females can adaptively adjust phenotype of their offspring via deposition of various compounds into eggs, including androgens and other hormones. Here, I investigated how egg yolk androgens (testosterone and androstenedione) related to environmental conditions and parental traits in the great tit (Parus major) across three breeding seasons. Male and female traits studied included age, condition and multiple feather ornaments, both carotenoid- and melanin-based (carotenoid and UV chroma of yellow breast feathers, area of black breast band and white cheek immaculateness). Yolk mass increased with laying temperature, laying date and area of male black breast band. Concentration of androgens increased with breeding density, territory quality and carotenoid chroma of male yellow breast feathers and was higher in mates of 1 year old as compared to older males. Yolk androgens were not related to any of the female traits analysed. These patterns were thus consistent with (1) social and environmental effects on yolk mass and composition and (2) both positive and negative differential allocation strategies of resource allocation in females. Overall, male traits were the most important predictors of egg yolk characteristics in this socially monogamous songbird.     

Effects of environmental manipulation on the locomotor activity and agonistic behavior of cultured juvenile American lobsters,Homarus americanus

The locomotor activity patterns and agonistic encounters of cultured juvenile (IX–XIII stage) American lobsters, Homarus americanus (Milne-Edwards) held in 8 experimental environments were investigated to examine their relative contribution to aggressive level. Three variables: (1) lobsters individually or communally maintained, (2) shelters present or absent, and (3) open or closed seawater system were tested in a set of factorial experiments. Locomotor activity of lobsters held in a 12 h: 12 h light: dark regime was recorded for individually separated lobsters for the first 5 days and for communally held lobsters for the next 5 days of a 10-day experimental period. All lobsters maintained individually had similar levels of activity. When the same individuals were communally maintained, there appeared to be social inhibition of activity. In addition, communally held lobsters without shelter were twice as active as those provided with shelter. The activity levels of groups of communally held lobsters with shelter progressively decreased over the experimental period. All lobsters became somewhat entrained to the light: dark cycle and were nocturnally active. However, the degree of entrainment was strongly influenced (P<0.001) by the presence or absence of shelter and other lobsters. Individually maintained lobsters all showed good entrainment, although those provided with shelter were slightly better entrained. The presence of conspecifics desynchronized activity patterns when shelter was not provided. Aggressive level, as measured by the frequency of agonistic encounters per 15-min observation period and the number and types of displays per encounter, decreased over an 8-day period for groups of 3 lobsters in all experimental environments. Lobsters maintained in a closed system initially showed higher frequencies of agonistic encounters than those held in an open system. The observed decrease in aggressive level with time primarily resulted from decreased activity levels. When two lobsters met, the probability of an agonistic encounter was about 0.8 throughout the experimental period, but decreasing activity levels resulted in progressively fewer encounters. The results suggest several approaches for limiting aggression among communally maintained lobsters in culture systems by environmental manipulation.     

Some relationships between size,food availability and energy balance in the ribbed musselAulacomya ater

The effects of body size and suspension density on filtration rates, assimilation efficiencies and respiration rates in the ribbed musselAulacomya ater (Molina) have been determined by means of short-term laboratory experiments. Filtration rates accelerate rapidly in response to increasing algal concentration up to approximately 10×10⁶ cellsDunaliella primolecta l^-1, beyond which a plateau is approached. Percentage increments are greatest in small individuals. Assimilation efficiencies are independent of body size, but decline rapidly with increasing ration to approach zero above 32×10⁶ cells l^-1. Increases in respiration rate accompany increments in filtration rate in all but the smallest size class tested. Filtration, assimilation efficiency and respiration measurements are used to calculate ingestion rations, assimilation rations and scope for growth for mussels of different sizes over a range of algal concentrations. Scope for growth, expressed as percentage change in body energy per day, is a declining function of body size, but larger individuals achieve their maximum growth rates at lower ration levels than smaller ones. Growth efficiency is independent of body size, and is maximal at 5×10⁶ cells l^-1, where 29 to 43% of ingested ration is converted into body energy. The applicability of these experimental results to natural ecosystems is discussed.     

Scent mediation of intraspecific,interspecific, and intergeneric agonistic behavior among sympatric species of voles (Microtinae)

Summary The hypothesis that the posterolateral scent glands of male microtine rodents (Muridae) direct the attacks of other males was tested.Montane voles (Microtus montanus) and sagebrush voles (Lagurus curtatus) were established as residents in terraria, and conspecific and heterospecific males, from which posterolateral scent glands had been unilaterally removed, were introduced. More attacks by residents were aimed at the glandular sides than at the glandless sides. Male M. montanus also attacked the unilateral hip gland of longtail voles (M. longicaudus), the growth of which had been induced by exogenous hormone inasmuch as this species does not normally possess these glands. Resident male M. longicaudus did not similarly attack the induced glands of conspecific males.The findings are related to demographic patterns, mterspecific relationships, and the evolutionary development of posterolateral scent glands among microtine rodents.