Responses of territorial and floater male Red‐winged Blackbirds to models of receptive females

Red‐winged Blackbirds (RWBL; Agelaius phoeniceus) have a polygynous mating system and, because territorial males commonly have harems of two to five females, some second‐year (SY) and after‐second‐year (ASY) males do not establish nesting territories, but become floaters. Previous studies have revealed high rates of extra‐pair copulations in this species and that sexually mature male floaters and territory owners do not differ in size, testosterone levels, or reproductive capability, suggesting that floaters may occasionally gain paternity. During May and June 2008, we observed the behavioral responses of floater males to taxidermic mounts (models) of female RWBL placed in a precopulatory position. Floaters intruded into territories during 46% of model presentations, with 20% of intrusions by ASY floaters and 80% by SY floaters. During intrusions, ASY floaters attempted to copulate with models 93% of the time compared to 80% for SY floaters. Copulations were successful during 30% of attempts by ASY males and 25% of attempts by SY floaters. The frequency of intrusions by ASY and SY floaters, attempted copulations by SY floaters, and successful copulations by ASY floaters increased as territorial males spent more time off their territories. Responses of floater males toward models in our study suggest that floater male RWBL attempt to exploit available breeding opportunities. The lack of evidence for extrapair young (EPY) fathered by floater male RWBL in previous studies, combined with our results indicating that the presence of territorial males limits floater intrusions, copulation attempts, and successful copulations, suggests that the reproductive success of floater males is limited in part by the aggressive behavior of territorial males.     

Behavioral syndromes influence mating systems: floater pairs of a lizard have heavier offspring

Variation in correlated behaviors or behavioral syndromes couldhave interesting effects on mating systems, especially if thevariation in syndrome exists in both sexes. Both males and femalesof the lizard Eulamprus heatwolei display two behavioral typesof a behavioral syndrome, defined by correlations between territorial,exploratory, and predator avoidance behaviors. We tested howthis variation in behavioral syndrome influences reproductivesuccess, pairing patterns, and offspring weight. We used spatialbehavior and residency in the field to identify territorialand floater individuals. Females were relocated to the laboratoryto give birth, and all offspring, dams, and potential sireswere genotyped to determine offspring paternity. During fieldsurveys, 164 lizards were caught of which 27.5% were territorialand the rest were floaters. Paternity was assigned to 66% ofthe 104 offspring produced by 33 dams. Territorial sires fathereda greater proportion of the offspring of territorial dams thanfloater sires. Larger territorial males were more likely tosire the entire clutch or share paternity with fewer additionalsires than smaller territorial males. Floater sire size, however,did not influence the number of fathers per litter. Floaterfemales produced heavier offspring than their territorial counterparts,and offspring fathered by floaters were heavier than maternalhalf-sibs fathered by territorial males. We speculate that differencesin offspring weight may be the result of differences in yolkprovisioning by females and parent genetic compatibility.     

Delayed plumage maturation in green‐backed flycatchers (Ficedula elisae): An evidence of female mimicry

Delayed plumage maturation (DPM) is the delayed acquisition of an adult color and pattern of plumage until after the first potential breeding period. Among the hypotheses proposed to explain DPM, the female mimicry hypothesis (FMH) has received considerable attention. FMH predicts that after‐second‐calendar‐year (ASY) males should attack ASY males more than second‐calendar‐year (SY) males and females, while no difference between the two latter. Few studies have been thought as support for FMH, while in fact most of them give no conclusive evidence to the assumption of FMH that ASY males are unable to distinguish SY males from females. Thus, other support besides behavioral experiments, such as it is physiologically impossible to discriminate SY males and females from avian perspective, is important in testing FMH. We studied color differences in six plumage patches between ASY male, SY male and female by analyzing reflectance spectra in the avian visual system, and tested the prediction of FMH by conducting intrusion experiments on territorial males with three kinds of conspecifics (i.e., ASY males, SY males and females) as territory “intruders” in the green‐backed flycatcher (Ficedula elisae). It is physiologically impossible for ASY males to distinguish SY males and females through the plumage color of crown, mantle, rump, and throat patches, moreover difficult to distinguish through breast and belly patches even under reasonable viewing conditions. ASY males were more aggressive toward ASY males than SY males and females with no differences between the two latter. SY males showed a slightly but not significantly higher attack intensity on ASY males than SY males and females. Our results suggest that female mimicry is more likely to be the explanation for DPM in the green‐backed flycatcher. To our knowledge, this is the first study combining avian visual system and behavioral experiments in testing hypotheses of DPM.     

Floater interference reflects territory quality in the Spanish Imperial Eagle Aquila adalberti: a test of a density‐dependent mechanism

We report on an 11‐year study of floater interference in a population of Spanish Imperial Eagles Aquila adalberti. We analysed changes over the years in the productivity of 15 territories to test predictions of two hypotheses of density‐dependent productivity in relation to the presence of floaters (birds without territories). According to the ‘interference' hypothesis, the frequency of intrusion by floaters increases with density, resulting in a decrease in productivity. Thus, in a high‐density population a negative relationship between floater intrusions and productivity of the territory is expected. In contrast, under the ‘habitat heterogeneity' hypothesis, as density increases a higher proportion of individuals is forced to occupy lower quality habitats. Support of this hypothesis requires that floaters detect differences in quality among territories and preferentially visit the better quality territories. Consequently, a positive relationship between floater intrusions and productivity is expected. Results showed that floaters tended to visit their natal area at the beginning of the breeding season. Among floater Eagles, males made significantly more intrusions per day than did females, but females stayed in the natal population for longer each year than males. Floater intrusions and productivity were highly positively correlated, supporting the ‘habitat heterogeneity' hypothesis; individuals were apparently able to assess the quality of a territory and, at the frequencies observed, their interference with the breeding pair had no obvious negative effect on productivity.     

Effect of alarm calling by male Red‐winged Blackbirds on nestling begging and female provisioning behavior

ABSTRACT Nestling begging and parental provisioning can attract nest predators and reduce reproductive success, so parents and their offspring might be expected to respond adaptively by minimizing predator‐attracting cues when predators threaten nests. Male Red‐winged Blackbirds (Agelaius phoeniceus) are well known for their antipredator alarm calls that contain information about the approach of potential nest predators. We examined the begging behavior of nestlings and the provisioning behavior of females in response to antipredator alarm calls of males to test the adaptive response hypothesis. Playback experiments provided no evidence that alarm calls function to switch off vocal begging; nestlings were equally likely to beg vocally during playback and control periods. Video recordings showed that male alarm calling had no significant effect on inappropriate vocal begging (in the absence of an adult), but significantly reduced the incidence of spontaneous calling (in the absence of begging). Adult females responded to male antipredator alarm calls by delaying their provisioning visits. In addition, although having no significant effect on use of nest‐arriving calls by females, male alarm calling significantly reduced their use of nest‐leaving calls. We conclude that nestling and female Red‐winged Blackbirds respond to male alarm calls in ways that might reduce the risk of predation, but nestlings beg vocally when females arrive to feed them, regardless of male alarm calling, perhaps to avoid a competitive disadvantage with broodmates.     

Territory quality affects the relative importance of habitat heterogeneity and interference competition in a long‐lived territorial songbird

Density‐dependent reproduction is commonly explained by either the habitat heterogeneity (HHH) or individual adjustment (IAH) hypothesis. Under the HHH, high quality territories are assumed to be occupied first. At higher density, occupation of low‐quality territories increases due to lower availability of high‐quality territories, which reduces mean reproductive success. Alternatively, the IAH assumes that increased competition at higher densities reduces reproductive success in all territories. For birds of prey, HHH plays an important role in territorial species, and IAH in socially breeding species. To test the generality of this hypothesis, we studied the mechanism behind density dependence in raven Corvus corax, a long‐lived passerine bird, using long‐term population data from a large number of territories. Population density decreased reproduction, which was explained by increased usage of low quality territories at higher density, supporting the HHH. Density reduced reproduction in low quality territories, but not in high and intermediate quality territories. We additionally compared the explanatory power of different models describing brood size, representing IAH, HHH, or a combination of both. The best model represented a combination of both hypotheses, in which the effect of density depended on territory quality. Our conclusion that both IAH and HHH are supported can be explained by the biology of ravens, where territorial adults not only experience interference competition with other territorial adults, but also with social groups of juveniles and floaters. We conclude that the relative importance of IAH and HHH may depend on variation in territory quality and social structure.     

Comprehensive paternity assignment: genotype,spatial location and social status in song sparrows,Melospiza Melodia

Comprehensive, accurate paternity assignment is critical to answering numerous questions in evolutionary ecology. Yet, most studies of species with extra‐pair paternity (EPP) fail to assign sires to all offspring. Common limitations include incomplete and biased sampling of offspring and males, particularly with respect to male location and social status, potentially biasing estimated patterns of paternity. Studies that achieve comprehensive sampling and paternity assignment are therefore required. Accordingly, we genotyped virtually all males and >99% of 6‐day‐old offspring over 16 years in a song sparrow (Melospiza melodia) population and used three complementary statistical methodologies to attempt complete paternity assignment for all 2207 offspring. Assignments were highly consistent across maximum likelihood methods that used solely genotype data, and heuristic and integrated Bayesian analyses that included data describing individual locations. Sires were assigned to >99% of all genotyped offspring with ≥95% confidence, revealing an EPP rate of c. 28%. Extra‐pair sires primarily occupied territories neighbouring their extra‐pair offspring; spatial location was therefore highly informative for paternity assignment. EPP was biased towards paired territorial males, although unpaired territorial and floater males sired c. 13% of extra‐pair offspring. Failing to sample and include unpaired males as candidate sires would therefore substantially reduce assignment rates. These analyses demonstrate the integration of genetic and ecological information to achieve comprehensive paternity assignment and direct biological insight, illustrate the potential biases that common forms of incomplete sampling could have on estimated patterns of EPP, and provide an essential basis for understanding the evolutionary causes and consequences of EPP.     

Reproductive Consequences of Aggression in a Territorial Songbird

Territorial aggression can influence males’ ability to obtain high‐quality resources and access to mates; however, in many species, the reproductive consequences of variation in aggression are unknown. In this study, we investigated how individual variation in aggressive behavior relates to reproductive success in socially monogamous, genetically polygynous song sparrows (Melospiza melodia). Prior research in this species shows that male song sparrows differ in their willingness to engage in agonistic interactions with territorial intruders and that individual variation in aggression appears to have functional significance. Aggressive males have been shown to obtain territories where females produce larger clutch sizes, suggesting that individuals who display high levels of territorial aggression are defending high‐quality territories or females. Further, aggressive males are considered a greater threat to territory‐holding males than less aggressive males. In this study, we ask whether individual differences in aggression are linked to differences in extra‐pair reproductive success, annual reproductive success, and offspring quality. We did not uncover a relationship between aggression and annual reproductive success or patterns of extra‐pair paternity. However, we found that the nestlings of aggressive males grew at a faster rate than the nestlings of less aggressive males. Future studies should attempt to identify mechanisms to explain the relationship between offspring growth rate and male aggression and investigate whether faster offspring growth rates translate to greater survival and recruitment of offspring.     

Estimation and limitation of numbers of floaters in a Eurasian Sparrowhawk population

Over a 20‐year period, the numbers of Eurasian Sparrowhawk Accipiter nisus nests found in a 200 km² area in south Scotland remained relatively stable (mean 33.3 pairs, CV = 10.6%). Nest numbers fluctuated from year to year in a manner expected of a population subject to density‐dependent regulation. The numbers of non‐breeders (floaters) could not be counted directly, but the number of female floaters was estimated, using known mortality rates, from the numbers of females recruited to the breeding population each year at different ages. Female floater numbers were estimated by two methods: Method A assumed that birds bred for the first time in their first, second or third year, in the same ratio as they were found breeding for the first time in the study area; and Method B assumed that all third‐year birds found breeding for the first time in the study area had bred previously, unknown to us, outside the area. Under Method A, floaters consisted of some 1‐year and some 2‐year birds; while under Method B, floaters consisted only of some 1‐year birds. Under both methods, the estimated number of female floaters fluctuated greatly from year to year, but under Method A they averaged 0.90 per female breeder, and under Method B they averaged 0.28 per female breeder. The Method B estimate was most consistent with other data and with the finding that some birds found breeding in the study area were likely to have bred previously outside the area (because of territory changes). Moreover, the mean and variance in female floater numbers estimated by Method B were similar in magnitude to the values obtained in a simulation model. In this model, breeding density was given a fixed ceiling, while breeding success was allowed to vary from year to year within the limits observed in the study area. It was concluded that (a) recruitment of floaters to the breeding sector was density dependent with respect to breeder numbers, i.e. broadly speaking, floaters filled gaps in the territorial system left by the deaths and movements of established breeders; and (b) floater numbers themselves were probably not regulated in a density dependent manner, but depended on whatever was the balance between annual additions (from reproduction and immigration) and subtractions (from mortality, emigration and entry to the breeding sector).     

Evidence of selection for egg crypsis in conspicuous nests

ABSTRACT The value of egg coloration as crypsis, once accepted as a general principle, has recently been questioned because most experiments have failed to show that egg coloration deters predation. The nest‐crypsis hypothesis postulates that, among species that build conspicuous nests, selection for egg crypsis is relaxed or absent because visually searching predators detect nests prior to eggs. I tested the nest‐crypsis hypothesis using the large, relatively conspicuous nests of American Robins (Turdus migratorius), and eggs that differed markedly in color that were collected from the nests of Red‐winged Blackbirds (Agelaius phoeniceus), Brewer's Blackbirds (Euphagus cyanocephalus), and Yellow‐headed Blackbirds (Xanthocephalus xanthocephalus). Each nest (N= 22) received a clutch of each species during three sequential predation trials that were 16 d in duration. The order of clutch presentation was randomized for each nest. Survival trends for Brewer's and Yellow‐headed Blackbirds were similar, and higher than those for clutches of Red‐winged Blackbirds. By the end of trials, overall survival of the three clutch types was roughly equivalent. However, clutches of Red‐winged Blackbird eggs, the most conspicuous egg type to the human eye, were discovered sooner by predators. Because the experimental design controlled for effects of nest crypsis, nest location, and nest size, this difference in egg survival can be attributed to differences in egg pigmentation. Thus, my results support a role for egg coloration as camouflage in conspicuous nests.     

Reproductive correlates of plumage coloration of female Mountain Bluebirds

Many studies have shown that the plumage coloration of male birds can act as an honest signal of quality, indicating benefits that a female could gain from pairing with a specific male. In some species, females also display ornamental plumage, but less is known about the function and potential adaptive significance of female coloration because most research has focused on male coloration. Male Mountain Bluebirds (Sialia currucoides) display full body, ultraviolet (UV)‐blue plumage, whereas female plumage is more subdued, with blue color focused on the rump, wing, and tail. During the 2011 and 2012 breeding seasons (May–July) near Kamloops, BC, Canada, we examined coloration of the rump and tail of female Mountain Bluebirds to determine if their plumage could act as an indicator of direct reproductive benefits (e.g., enhanced parental care or reproductive success) to potential mates. We found no relationship between female plumage coloration and either provisioning rate or fledging success. However, female coloration varied with age, with after‐second‐year (ASY) females having brighter, more UV‐blue tail feathers than second‐year (SY) females. In addition, ASY females with brighter, more UV‐blue tails had larger clutches. We also observed positive assortative mating by tarsus length. Because previous work with other species suggests that female body size may be a good predictor of breeding success, males could potentially benefit from pairing with larger females. However, reproductive success did not vary with female size in our study. Although our evidence that structural plumage coloration of female Mountain Bluebirds is a signal of direct reproductive benefits for males (e.g., higher reproductive success) is limited, our results (i.e., ASY females with brighter tails than SY females, and ASY females with brighter tails having larger clutches) do suggest the potential for sexual selection to act on female coloration.     

Mating tactics and male wing dimorphism in the damselfly,Mnais pruinosa costalis selys (Odonata: Calopterygidae)

Males of the damselfly,Mnais pruinosa costalis, exhibit wing color dimorphism: one form has orange wings, and the other hyaline wings which resemble female wings. The former is usually territorial and the latter uses sneaky mate securing tactics. When orange-winged males failed to establish territory, they became floaters that day. Hyaline-winged males perched around their territories and often, formed in tandem without any apparent courtship behavior when they found females. Their copulation frequency was higher and copulation duration longer than those of territorial males. A few females oviposited without remating. Total oviposition duration of females with which a hyaline-winged male mated was more than 32 min per male on average in a day Females that copulated with hyaline-winged males often remated with orange-winged residents before oviposition. Total duration of oviposition bouts of females after mating with floaters was short (15 min), while that with territorial residents was long (66 min). As a result, total oviposition duration of females with which an orange-winged male mated was about 40 min in a day. The reproductive success of the hyaline-winged males may be similar to that of the orange-winged males.     

Should I Stay or Should I Go Now: Late Establishment and Low Site Fidelity as Alternative Territorial Behaviors

In species in which males defend territories for breeding, males may differ in territorial behavior; alternative behaviors among territorial males are not well understood. In our long‐term study of partially‐migratory song sparrows, we have observed that most territorial males establish territories before females begin nesting and remain site‐faithful both within and between breeding seasons; however, some males establish territories later in the season (late establishers) and/or change territory locations either within or between seasons (movers). Whether late establishment or moving are equally successful strategies for territory defense, or best‐of‐bad‐job options, is not known. Here, we compare the frequencies of these behaviors to demographic variables over a 9‐yr period and compare lifetime tenure and early season nesting success for males who differ in site fidelity and timing of territory establishment. Across years, late establishing was negatively correlated with the return rate of previously territorial males; moving was positively correlated with the number of occupied territories at the start of the breeding season (territory density). While moving was independent of number of years on territory, late territory establishment only occurred in a male’s first year as a territory holder. Of 88 males, 25% established their first territory late, primarily in undefended space; 31% moved. Late and early establishers did not differ in lifetime tenure; movers, however, had longer lifetime tenure than site‐faithful males. Among early establishers, movers and non‐movers did not differ in the number of successful early nests/year or number of young fledged/year; among late establishers, however, movers had significantly higher early nesting success by both measures. Late establishers who moved had higher early season nesting success and higher early season nesting success/year than site‐faithful early establishers. Thus, individual variation in the timing of territory establishment and site fidelity may be facultative alternative territorial strategies.     

Optimal floating and queuing strategies: consequences for density dependence and habitat loss

Field studies of many vertebrates show that some individuals (floaters) do not defend territories even when there is space for them to do so. We show that the evolutionarily stable strategy (ESS) for the threshold territory quality at which floating takes place is that which maximizes the size of the floating population (but not the total population, breeding population, or reproductive output). The ESS is solved separately for two assumptions: whether individuals wait to occupy a single territory or multiple territories and whether queuing rules are strict or if all waiting individuals are equally likely to obtain the next territory. The four combinations of these assumptions all give the same evolutionarily stable population size of both floaters and breeders. At the ESS, only territories with expected lifetime reproductive success (LRS) exceeding 1 should be occupied, which introduces a limit to ideal habitat selection. The behavioral decision to float alters the shape of the density-dependent response, reduces the equilibrium population size, and affects the response of the population to habitat loss. Specifically, the floater: breeder ratio is directly related to average breeding habitat quality, and the floater population size will decrease more than the breeding population size if better than average quality habitat is lost.     

Social status influences microhabitat selection: breeder and floater Eagle Owls Bubo bubo use different post sites

Social status can be reflected in many aspects of an individual’s behaviour and ecology, including habitat use and conspecific interactions. In territorial species where at least two social groups – breeding birds and non‐territorial floaters – are recognized, the diverse tasks associated with territorial ownership can lead territory holders to behave differently from the non‐territorial part of the population. Territory holders defend their breeding area and reproduce, whereas floating individuals are dispersing and lead a more transient life, during which they do not show any territorial behaviour even when settling in a more or less fixed area (known as the stop phase). As social interactions are based on visual and vocal cues, the use of specific sites for sending and/or receiving signals can be a crucial choice in an animal’s life. By analysing the post‐site selection of Eagle Owl Bubo bubo breeders and floaters during their nocturnal activity, we found that: (1) territory holders selected more visible and dominant posts than non‐territorial floaters; (2) the choice of posts made by floating individuals did not differ between the wandering and stop phases of dispersal; and (3) floating females intruded more frequently than floating males within a breeder’s home‐range. These findings highlight the fact that two social strategies are possible within the same species, depending on an individual’s social status and its related tasks. Breeders could take advantage of visible locations to declare their status as territory holders, whereas floaters could benefit from a more secretive life to wander unnoticed among occupied territories. This secretive life would help floaters to reduce the risks associated with conspecific aggression. Finally, the greater occurrence of floating females within breeders’ home‐ranges can be explained by the fact that female incursions in a breeder’s home‐range are less risky than male intrusions.     

Uncertainty in territory quality affects the benefits of usurpation in a Mediterranean wrasse

Individuals should defend sites when the expected benefits ofthe territory exceed the cost of defense. However, if territoryqulaity is unpredictable or difficult to assess, the expectedpattern of territorial behavior is less clear. In a Mediterraneanwrasse, Symphodus ocellatus, mating success is skewed with 2%of nesting males getting more than 20% of the spawning success.Yet, variation in mating success is not explained by any knownphysical characteristic of males or their territories. Instead,females prefer nests with a recent history of mating successbecause males are less likely to desert the offspring she leavesbehind. Thus, territory quality is transient and determinedby interactions between the sexes. I measured the frequencyof territorial takeovers and the uncertainty in mating successamong days at a nest. Observations indicated that S. ocellatusmales usurped their neighbor's successful nests when males wereunsuccessful and larger than their successful neighbor. Sitesthat achieved mating success had a significantly higher probability(0.84) of remaining sucessful between consecutive days thanunsuccessful territories had of becoming successful (0.30).Unsuccessful males obtained higher and more certain fitnessreturns if they usurped a successful neighbor's territory. Interactionswithin and between the sexes drive uncertainty in success, whichinfluences territorial behavior in this species.     

Cautious response of inexperienced birds to conventional signal of stronger threat

Several studies demonstrated that bird song functions as a first line of territorial defence. The efficiency of deterring rivals depends strongly on the strategy of singing used (e.g. alternating/overlapping singing, singing with low/high rate, matching song type of a rival or singing different type). Causes of between males variation during countersinging are still not fully understood, especially when different signals have similar production costs and their meaning is assigned by arbitrary convention (conventional signalling). We tested whether an oscine bird with small repertoire size, the ortolan bunting Emberiza hortulana , differentiate strategy of responding to song of an intruder in relation to its age and threat value of signals. We performed playback experiments to measure response of second year (SY) and after second year (ASY) males to a song of low (eventual variety singing) and high (immediate variety singing) threat value. We found substantial differences in response to playback, which were related both to the type of stimuli used and age of responding males. Both SY and ASY males gave more calls than songs in response to immediate variety playback, which suggest stronger vocal response to the signal of higher threat value. Approaching loudspeaker was similar for both age classes when lower threat value signal was played back, while simultaneously SY males clearly avoided approaching loudspeaker when stronger threat values signal was played back. We conclude that ortolan bunting differentiate response to signal of different threat value and that the strength of response depends on the age of a male. This study provides experimental evidence that age of receiver affects its response to a territorial intruder. It also demonstrates that observed in many studies variation in response to playback may be an effect of age differences between males, which rarely is controlled.     

Seasonal and age-related changes in blood parasite prevalence in Dark-eyed Juncos (Junco hyemalis, Aves, Passeriformes)

We determined seasonal changes in blood parasite infections in a free-living population of Dark-eyed Juncos (Junco hyemalis) breeding in interior Alaska (65 degrees N; 148 degrees W). The common parasites found in blood smears were Leucocytozoon fringillinarum (56%), Trypanosoma avium (33%), and Haemoproteus fringillae (9%). In males, parasite prevalences were relatively high at arrival on breeding grounds and increased during the breeding season. Intensity of infection with Leucocytozoon also increased between spring and summer, and then decreased at the time of migration (September). This decrease did not occur in adult females. Elevated prevalences during the breeding season probably reflected the addition of new cases via vector activity to positive status resulting from spring relapse. We observed neither an association between parasite species nor a consistent relationship between parasite intensity and body condition. To further study relationships between reproductive system activity and parasite infections, we compared prevalences in adult males that were undergoing their first cycle of gonadal development and regression (males in their second calendar year, or SY) with those of older males (males in their third or more calendar year, i.e., after-second-year males or ASY). Circulating testosterone concentrations declined in both groups between arrival on breeding grounds (end of April-early May) and the end of the reproductive period (July), and they were higher in May in ASY than in SY males. At the peak of the breeding season (June), ASY males also had a higher parasite prevalence than SY males. This difference may have resulted from immunosuppressive effects of gonadal hormones and/or from behavioral differences between SY and ASY males such that older males were exposed to more insect vectors than younger males. .     

Effect of competitor abundance on feeding territoriality in a grazing fish,the ayuPlecoglossus altivelis

The grazing fish, ayu,Plecoglossus altivelis Temminck & Schlegel, establishes feeding territorialiry during the young stage. The population density fluctuates from year to year by more than a hundredfold, but the determinant of territory size is less well known. The feeding territoriality of ayu was examined under simulated habitat conditions where fish density was manipulated and food resources were renewable. Fish competed for algae attached to the substrata and were divided into residents with territories, and floaters without territories. By experimental alteration of fish density the number of residents increased with density and rerritory size decreased with density. Floaters intruded into territories in a school to feed on algae, which induced overt aggression of the resident and reduced the productivity of algae growing there. Both the intruding frequency of floaters over territorial areas and their feeding pressure on algae increased at higher floater density. Floaters functioned to shift cost-benefit relationships for various territory sizes. They acted as food competitors to restrict territory size below a maximum through competitive interference. Although the growth rate of residents was inversely related to fish density, residents grew faster than floaters in each group. Under a given set of competitor abundances, economic defensibility determined territory size.     

Sex roles, parental experience and reproductive success of eastern kingbirds, Tyrannus tyrannus

We quantified parental behaviour of eastern kingbirds during the incubation and nestling periods to determine parental roles, and to examine the impact of previous breeding experience (defined as having bred on the territory in the past) on behaviour and reproductive success. Females performed all incubation, while males spent more than 60% of their time in vigilant or nest guarding behaviour during incubation. Parental roles were not defined as sharply during the nestling period. Females spent more time vigilant, but males provisioned young at only 54% of the rate of females. Vigilance and nest watching were still primarily male duties. Male and female behaviour did not vary with the pair's combination of experience (e.g. experienced-experienced versus inexperienced-inexperienced in previous-current breeding season, respectively) during either phase of reproduction, but experienced males were more vigilant during incubation and fed young relatively more than inexperienced males. Experienced females were also more efficient foragers. Although behaviour did not differ among the four combinations of pair experience, inexperienced pairs none the less lost the most young to starvation and predation. Consequently, inexperienced pairs fledged one less nestling per nesting attempt than did pairs with at least one experienced breeder. Our results suggest that having at least one experienced breeder substantially improved a pair's reproductive success. We propose that female site fidelity is a safeguard to avoid the lower breeding success a female would incur if she were to move to a new territory and breed with an inexperienced male. Copyright 1999 The Association for the Study of Animal Behaviour.