Macroecological patterns of resource use in resident and migratory hummingbirds

Hummingbirds (Family Trochilidae) are key pollinators in several biodiversity hotspots, including the California Floristic Province in North America. Relatively little is known about how hummingbird diets change throughout the year, especially with regard to how migratory hummingbirds affect resident hummingbirds at stopover sites. In this study, we examine how hummingbird species, migratory status, sex, geographic region and local plant diversity influence floral resource use before, during, and after an influx of migratory hummingbirds (primarily Rufous hummingbirds, Selasphorus rufus) across California. We expected distinct floral resource use based upon species’ migratory status (resident vs. migrant), sex, sampling period, and geographic region. We employed DNA metabarcoding to detect plant DNA in hummingbird fecal samples to analyze diet diversity, composition, overlap, and interaction networks. We found significant effects of sex, sampling period, and migratory status on the alpha and beta diversity of plant taxa present in fecal samples. Analyses of Anna's hummingbirds (Calypte anna) alone revealed that female fecal samples contained higher plant species richness. In addition to hummingbird-pollinated plants, fecal samples also contained non-ornithophilous plants and species of agricultural importance. Diet overlap and plant-pollinator network analyses revealed high overlap in plant taxa used between hummingbird species, and networks were more connected, less nested, and less specialized than null models. DNA metabarcoding is minimally invasive and provides a detailed view of hummingbird diet, permitting large-scale studies. Insights into hummingbird diets are especially valuable given the logistical difficulties of directly observing floral visitation and foraging across broad temporal and spatial scales.     

The role of size and dominance in the feeding behaviour of coexisting hummingbirds

Interspecific competition can strongly influence community structure and limit the distribution and abundance of species. One of the main factors that determine hummingbird community structure is competition for food. The temporal and spatial distribution of nectar has a strong impact on hummingbird assemblages, shaping foraging niches according to hummingbird dominance and foraging strategy. We investigated whether body size and the degree of aggressive dominance influence feeding behaviour of hummingbirds in a temperate forest in northwestern Mexico (El Palmito, Mexico) when winter migrant hummingbirds are present in the community. First, we determined the dominance status of hummingbirds and evaluated the relationship between dominance and body mass, wing disc loading and migratory status. Secondly, we determined how hummingbird species used plant species differently. Thirdly, we examined whether the most dominant hummingbird species defended floral patches with more energy and/or with a larger number of flowers. At each flower patch, hummingbird species, number of hummingbird interactions, feeding time and number of flowers present were recorded. The total number of calories available within each floral patch was also determined. Our results demonstrate that the dominance hierarchy of 13 hummingbird species (migratory and resident) was correlated with body size but not wing disc loading, and that members of the hummingbird community showed a clear separation in resource use (by plant species). Hummingbirds at the top of the dominance hierarchy defended and fed on the best flower patches, defined by the quantity of calories available. Hence, the feeding behaviour of hummingbirds at El Palmito depends on the abundance of plant species used by hummingbirds and on the amount of energy available from each flower patch. Thus, hummingbird body size, aggressive dominance and defence of quality flower patches determines niche partitioning among species.     

Effects of lamb rearing conditions on aggression and dominance relationships

The effects of rearing conditions on agonistic behavior were studied using lambs reared: (1) in a flock; (2) individually with a ewe; (3) with other lambs; (4) as individuals. The lambs were paired in a noncompetitive situation preweaning, and again postweaning, after 10 weeks of cohabitation in one group; competitive paired tests were also conducted postweaning. During the preweaning paired tests, the socially deprived lambs exhibited play and investigatory behavior while the flock reared lambs ignored the other lambs or were aggressive. Neither the postweaning noncompetitive nor the competitive paired tests revealed disparity in behavior among treatments.     

Temporal partitioning of a floral resource by territorial hummingbirds

Most studies of territoriality in hummingbirds have focused on intraspecific competition for resources and the consequences for the spatial distribution of individuals within a habitat. As a result, we know little of the effects of interspecific competition for resources and less still of temporal resource partitioning. Here I describe the interactions of four species of tropical hummingbird which defended the same territory at different stages in the flowering period and at different times of the day. The pattern of territory defence was greatly influenced by the dominance hierarchy between species and the costs and benefits of territory ownership. I used a simple economic model to calculate the predicted territory size based on four potential strategies. Hummingbirds appeared to be defending territories of the smallest economical size, agreeing with two hypotheses: (1) that hummingbirds minimize the cost of territory ownership and (2) that hummingbirds maximize the time spent sitting. The model predicted accurately the observed pattern of territory acquisition; hummingbirds initiated defence as soon as the territory contained sufficient resources and were either displaced by a larger species or replaced by a smaller one as the value of the territory changed.     

Parasites, aggression and dominance in male upland bullies

The effects of the digenean trematode parasite Telogaster opisthorchis on aggression and competitive ability in male upland bullies ( Gobiomorphus breviceps ) were examined using mirror-image stimulation (MIS) techniques and dyadic contests for nest sites between pairs of males. Parasite load had no significant effect on male aggression or success in dyadic contests, nor was aggression a predictor of the likely winner of these contests. The results are discussed with relation to the role of parasites in male-male competition and to possible problems in using MIS techniques for studies on dominance and aggression in fish.     

Influence of social dominance rank on diet quality of pronghorn females

The relationship between social dominance rank and diet qualitywas examined in pronghorn (Antilocapra americana) females atthe National Bison Range in Montana. Rank was ascertained byobservation of agonistic interactions between study animals.Diet quality of the same individuals was quantified by measuringthe concentration of the chemical indicator of forage quality,2, 6 diaminopimelic acid (DAPA) present in feces. A significant, quadratic relationship was found in which high-ranked and low-rankedfemales showed high levels of DAPA, and middle-ranked femalesshowed low levels of DAPA. This finding indicated that highand low-ranked females acquired better quality diets than middle-ranked females. I discuss possible explanations for this interestingresult which, to my knowledge, has not been anticipated in the literature.     

A model of dominance and resource division among a group of animals of differing quality

Individuals of many species spend a large portion of their lives within groups of conspecifics. Within such groups, there can be considerable diversity in size and other traits, with some, perhaps larger or stronger, animals laying claim to a greater than fair share of available resources. We use the word quality as a single measure of individual animal phenotypes within such a group and develop a model of resource division through contests between animals of differing quality. We investigated the effect of varying environmental factors on the level of aggressiveness used in such contests and the division of resources among the group. A numerical example shows that maximum discriminations between individuals on the basis of quality occur for medium-sized costs of aggressive encounters, but resources are divided more equitably at either extreme, when costs are either high or low. Received: April 9, 2001 / Accepted: September 16, 2001     

Red clothing increases perceived dominance,aggression and anger

The presence and intensity of red coloration correlate with male dominance and testosterone in a variety of animal species, and even artificial red stimuli can influence dominance interactions. In humans, red stimuli are perceived as more threatening and dominant than other colours, and wearing red increases the probability of winning sporting contests. We investigated whether red clothing biases the perception of aggression and dominance outside of competitive settings, and whether red influences decoding of emotional expressions. Participants rated digitally manipulated images of men for aggression and dominance and categorized the emotional state of these stimuli. Men were rated as more aggressive and more dominant when presented in red than when presented in either blue or grey. The effect on perceived aggression was found for male and female raters, but only male raters were sensitive to red as a signal of dominance. In a categorization test, images were significantly more often categorized as ‘angry’ when presented in the red condition, demonstrating that colour stimuli affect perceptions of emotions. This suggests that the colour red may be a cue used to predict propensity for dominance and aggression in human males.     

Stress coping style predicts aggression and social dominance in rainbow trout

Social stress is frequently used as a model for studying the neuroendocrine mechanisms underlying stress-induced behavioral inhibition, depression, and fear conditioning. It has previously been shown that social subordination may result in increased glucocorticoid release and changes in brain signaling systems. However, it is still an open question which neuroendocrine and behavioral differences are causes, and which are consequences of social status. Using juvenile rainbow trout of similar size and with no apparent differences in social history, we demonstrate that the ability to win fights for social dominance can be predicted from the duration of a behavioral response to stress, in this case appetite inhibition after transfer to a new environment. Moreover, stress responsiveness in terms of confinement-induced changes in plasma cortisol was negatively correlated to aggressive behavior. Fish that exhibited lower cortisol responses to a standardized confinement test were markedly more aggressive when being placed in a dominant social position later in the study. These findings support the view that distinct behavioral-physiological stress coping styles are present in teleost fish, and these coping characteristics influence both social rank and levels of aggression.     

Relationship of chin spot size to dominance in the black-chinned mouthbrooding cichlid fish (Sarotherodon melanotheron)

Aggressive interactions of male Sarotherodon melanotheron were studied to determine the communicative value of chin spot size in relation to dominance. In addition, the effects of residency and size on the aggressiveness of this fish were determined. Two-factor analysis of variance was used for frequencies of each modal action pattern for residency and size. Results show that residency played a major role in the outcome of an aggressive interaction, whereas size had little effect. Dominance of each experimental fish was calculated using Barlow & Ballin's (1976) dominance index. A chin spot ratio was obtained by dividing the chin spot area by the total surface area of the individual fish. Simple linear regression was conducted to determine if dominance and chin spot size were correlated and a positive linear relationship was found to exist between the two variables.     

Control of aggression and dominance in white-throated sparrows by testosterone and its metabolites

In three experiments, we investigated whether testosterone itself or its metabolites activate aggression and dominance in white-throated sparrows Zonotrichia albicollis. Groups of five to six sparrows, each treated with a different steroid implanted subcutaneously, were observed in outdoor aviaries during late winter to determine the birds' rates of aggression (supplantations and attacks scaled to the number of available subordinates) and dominance rankings with opponents not previously encountered. In Experiment 1, testosterone (T) had a greater effect on aggression and dominance than did androstenedione, 5α-dihydrotestosterone (D), androsterone, or estradiol (E). In Experiment 2, birds with T or D + E had higher aggression scores and dominance ranks than birds with either D or E alone. Birds with T and D + E did not differ. The testosterone metabolites, D and E, thus acted synergistically to determine rates of aggression and dominance ranks. To corroborate these results, in Experiment 3 we treated T-implanted birds with the following blocking agents: ATD, expected to reduce conversion of T to E (AT birds); progesterone, expected to reduce conversion of T to D (PT birds); or both (APT birds). The APT birds had lower aggression scores and dominance ranks than did AT or PT birds, despite having higher mean levels of circulating T than AT or PT birds or birds implanted with T alone. Cyproterone acetate also reduced aggression scores and dominance in T-implanted birds. We conclude that the hormonal control of aggression and dominance in these birds requires conversion of testosterone to both androgenic and estrogenic metabolites.     

Body mass as a supertrait linked to abundance and behavioral dominance in hummingbirds: A phylogenetic approach

Body mass has been considered one of the most critical organismal traits, and its role in many ecological processes has been widely studied. In hummingbirds, body mass has been linked to ecological features such as foraging performance, metabolic rates, and cost of flying, among others. We used an evolutionary approach to test whether body mass is a good predictor of two of the main ecological features of hummingbirds: their abundances and behavioral dominance. To determine whether a species was abundant and/or behaviorally dominant, we used information from the literature on 249 hummingbird species. For abundance, we classified a species as “plentiful” if it was described as the most abundant species in at least part of its geographic distribution, while we deemed a species to be “behaviorally dominant” when it was described as pugnacious (notably aggressive). We found that plentiful hummingbird species had intermediate body masses and were more phylogenetically related to each other than expected by chance. Conversely, behaviorally dominant species tended to have larger body masses and showed a random pattern of distribution in the phylogeny. Additionally, small‐bodied hummingbird species were not considered plentiful by our definition and did not exhibit behavioral dominance. These results suggest a link between body mass, abundance, and behavioral dominance in hummingbirds. Our findings indicate the existence of a body mass range associated with the capacity of hummingbird species to be plentiful, behaviorally dominant, or to show both traits. The mechanisms behind these relationships are still unclear; however, our results provide support for the hypothesis that body mass is a supertrait that explains abundance and behavioral dominance in hummingbirds.     

Influence of slaughter age and carcass suspension on meat quality in Angus heifers

This study investigated the effects of pelvic suspension and slaughter age on longissimus thoracis et lumborum (LTL) from 40 heifers with at least 75% Angus breeding. A total of 20 heifers were slaughtered directly from pasture at 18 months of age, and carcass sides were hung either by the Achilles tendon or the pelvic bone. The other 20 heifers were assigned to an additional winter housing period and slaughtered at 22 months of age; carcass sides were hung only by Achilles suspension. All carcasses were electrically stimulated and assessed according to the EUROP carcass classification system. In addition, the LTL muscles were aged for 7 or 14 days before meat quality was evaluated for intramuscular fat (IMF), drip loss, colour, shear force, compression and sensory analysis. The 22-month-old heifers were heavier, fatter and had more IMF than 18-month-old heifers. Conformation scores (muscling) did not differ between the two slaughter groups. Pelvic suspension reduced both between- and within-animal variation for peak force, total energy and compression peak force. For the 18-month-old heifers, pelvic suspension also decreased peak force, total energy and compression variables for the LTL muscles from both ageing periods, whereas Achilles-suspended samples had lower shear force values only at day 14. Sensory analysis showed that pelvic-suspended sides had greater tenderness, lower bite resistance, less threadiness, higher juiciness and meat flavour and less visible marbling than meat from Achilles-suspended sides. Pelvic-suspended sides at 18 months of age were similar in peak force and total energy values to the 22-month-old heifers. The importance of ageing the Achilles-suspended sides was more obvious for samples from 18-month-old heifers than from the 22-month-old animals. The correlations between the different instrumental measurements and sensory tenderness were considerably higher for carcasses suspended by the Achilles tendon (r = −0.55 to 0.20) than for those hung by the pelvic bone (r = −0.25 to 0.19). More correlations between sensory-evaluated tenderness and shear variables were significant after 7 days (n = 6) of ageing than after 14 days (n = 4) of ageing. This study clearly shows the benefits of pelvic suspension, which reduces the need for additional feeding after pasture.     

The relationships among testosterone, conditioned aggression, and dominance in male pigeons

Pigeons lowest in their homeloft dominance hierarchies were either injected with testosterone, conditioned to peck the more dominant birds, or both, and were then returned to their homelofts. Frequency of pecks and threats received by each animal was recorded. Testosterone alone had no effect. Conditioned birds were pecked significantly fewer times after treatment. The combination of hormone treatment and conditioning produced the greatest enhancement in dominance. Differences in the patterns of fighting behavior are noted and results are discussed in a discrimination learning framework as an interaction between androgen and learned responses.     

Testing a simple rule for dominance in resource competition

Competition for limiting resources long has been considered an important factor generating community structure. A minimal model of resource competition predicts that the species that reduces the limiting resource R to the lowest level ([Formula: see text]) will exclude its competitors. Whether this "[Formula: see text] rule" is robust to violations of model assumptions remains largely unknown. I conducted a competition experiment with four species of bacterivorous protists in laboratory microcosms and predicted the outcome from each species' [Formula: see text] value. I also examined how the outcome of competition, species abundances, and the effect of protists on bacterial density varied with productivity. Microcosms were unstirred batch cultures containing a variety of bacteria, challenging the robustness of the simplest competition models. Protists with low [Formula: see text] values were less affected by competition, although competing protists often coexisted. The values of [Formula: see text] can predict competitive dominance, even in the absence of competitive exclusion. Other model predictions were less robust. Contrary to expectation, densities of grazed bacteria increased with productivity, and the effect of some protists on bacterial density did not vary with productivity. Bacterial heterogeneity may account for deviations from model predictions. Further experiments should examine the conditions under which simple rules can be expected to identify dominant species.