Variance‐Sensitive Green Woodhoopoes: A New Explanation for Sex Differences in Foraging?

Studies of cooperatively breeding birds rarely benefit from the extensive research on adaptive foraging behaviour, despite the potential for concepts such as state‐dependent foraging to explain many aspects of behaviour in social groups. For example, sex differences in preferred foraging techniques used by green woodhoopoes, Phoeniculus purpureus, have previously been explained by sexual dimorphism in bill length and the benefits afforded by foraging specialization and niche differentiation within cooperative groups. Contrary to this argument, there were no sex differences in mean foraging success and/or prey size captured when males and females used the same foraging techniques. Subordinates of both sexes did experience lower and more varied foraging success compared with dominants, but probably only as a consequence of competition or inexperience. However, dominant males experienced greater variance in individual foraging success compared with dominant females, and dominant males also experienced greater variances in prey size when using their preferred foraging techniques. Dominant males therefore appeared to specialize in foraging techniques that provided more variable rewards, whilst dominant females consistently chose to minimize variation in reward. Dominant females also experienced less variance in foraging returns when using the same techniques as males, suggesting a possible link with sexual dimorphism in bill length. Partitioning of foraging niches in dominant green woodhoopoes therefore appears to be better explained by sex differences in variance (risk) sensitivity to foraging rewards. We suggest that this kind of detailed analysis of state‐dependent foraging has the potential to explain many of the crucial age and sex differences in behaviour within cooperative groups.     

Cue use affects resource subdivision among three coexisting hummingbird species

Competition for food can influence the coexistence of speciesvia habitat selection, and learned behavior can influence foragingdecisions. I investigated whether learned behavior and competitionact together to influence species interactions between threecoexisting hummingbird species: black-chinned (Archilochusalexandri), blue-throated (Lampornis clemenciae), and magnificent(Eugenes fulgens) hummingbirds. I found that color cue useby individuals affects not only their foraging choices butalso population-level responses to competition. I presented hummingbirds two types of habitats (rich and poor feeders).All birds shared a preference for the rich feeders, but shiftedpreference toward poor feeders in response to competition.I used color cues to manipulate the amount of information availableto birds and examined the effects of two information states(complete or incomplete) on their foraging choices. I examined hummingbirds' preferences for the rich feeders when both competitordensities and information varied. To relate foraging choicesto energetic intake, I also analyzed energy gained during asingle foraging bout. Males of all species exhibited strongpreferences for rich feeders when they foraged with complete information and low competitor densities. Without complete information,the two subordinate species (black-chinned and magnificent)shifted preference away from rich feeders in response to highdensities of the dominant species (blue-throated). Each subordinatespecies shifted in a unique way: black-chinned hummingbirdsreduced foraging efficiency, while magnificent hummingbirdsreduced foraging time. Birds foraging with complete information remained selective on rich feeders even at high competitor densities.Thus, learned information affected competitive interactions(for rich feeders) among these species.     

Hummingbird responses to gender-biased nectar production: are nectar biases maintained by natural or sexual selection?

Pollinators mediate the evolution of secondary floral traits through both natural and sexual selection. Gender-biased nectar, for example, could be maintained by one or both, depending on the interactions between plants and pollinators. Here, I investigate pollinator responses to gender-biased nectar using the dichogamous herb Chrysothemis friedrichsthaliana (Gesneriaceae) which produces more nectar during the male floral phase. Previous research showed that the hummingbird pollinator Phaethornis striigularis visited male-phase flowers more often than female-phase flowers, and multiple visits benefited male more than female fecundity. If sexual selection maintains male-biased rewards, hummingbirds should prefer more-rewarding flowers independent of floral gender. If, however, differential rewards are partially maintained through natural selection, hummingbirds should respond to asymmetry with visits that reduce geitonogamy, i.e. selfing and pollen discounting. In plants with male biases, these visit types include single-flower visits and movements from low to high rewards. To test these predictions, I manipulated nectar asymmetry between pairs of real or artificial flowers on plants and recorded foraging behaviour. I also assessed maternal costs of selfing using hand pollinations. For plants with real flowers, hummingbirds preferred more-rewarding flowers and male-phase morphology, the latter possibly owing to previous experience. At artificial arrays, hummingbirds responded to extreme reward asymmetry with increased single-flower visits; however, they moved from high to low rewards more often than low to high. Finally, selfed flowers did not produce inferior seeds. In summary, sexual selection, more so than geitonogamy avoidance, maintains nectar biases in C. friedrichsthaliana, in one of the clearest examples of sexual selection in plants, to date.     

Reversal Learning and Risk‐Averse Foraging Behavior in the Monarch Butterfly,Danaus plexippus (Lepidoptera: Nymphalidae)

Learning ability allows insects to respond to a variable environment, and to adjust their behaviors in response to positive or negative experiences. Pollinating insects readily learn to associate floral characteristics, such as color, shape, or pattern, with appetitive stimuli, such as the presence of a nectar reward. However, in nature pollinators may also encounter flowers that contain distasteful or toxic nectar, or offer highly variable nectar volumes, providing opportunities for aversive learning or risk‐averse foraging behavior. Whereas some bees learn to avoid flowers with unpalatable or unreliable nectar rewards, little is known about how Lepidoptera respond to such stimuli. We used a reversal learning paradigm to establish that monarch butterflies learn to discriminate against colored artificial flowers that contain salt solution, decreasing both number of probes and probing time on flowers of a preferred color and altogether avoiding artificial flowers of a non‐preferred color. In addition, when we offered butterflies artificial flowers of two different colors, both of which contained the same mean nectar volume but which differed in variance, the monarchs exhibited risk‐averse foraging: they probed the constant flowers significantly more than the variable ones, regardless of flower color or butterfly sex. Our results add to our understanding of butterfly foraging behavior, as they demonstrate that monarchs can respond to aversive as well as appetitive stimuli, and can also adjust their foraging behavior to avoid floral resources with high variance rewards.     

An empirical demonstration of risk-sensitive foraging preferences

We report laboratory experiments with yellow-eyed juncos (Junco phaeonotus) revealing that the birds' foraging preferences for variable rewards respond not only to the mean, but also the variance, of food rewards. The nature of their preferences for variable rewards is related to their expected daily energy budget. We summarize the birds' preferences in utility functions for energetic rewards. Since mean reward size is inadequate to predict their behaviour, we believe that foraging models should consider environmental stochasticity and an animal's response to this variation.     

An Assemblage of Hummingbird-pollinated Flowers in a Montane Forest in Southeastern Brazil

Relationships between ornithophilous flowers and hummingbirds have been little studied in southern South America, where hummingbird species richness is low. We studied an ornithophilous flower assemblage and the hummingbird pollinators in a montane forest in southeastern Brazil. Twenty-three native hummingbird-pollinated plant species in 21 genera and 14 families were observed. Bromeliaceae, Fabaceae, Gesneriaceae, and Lobeliaceae are represented by more than one species within the assemblage. Flower shapes vary from narrow tube to bowl-shape, but tubular flowers prevail. The variety of flower shapes and sizes results in diverse pollen placement on the body parts of hummingbird visitors, although pollen is deposited mostly on the bill. Sugar concentration in nectar averages 22.1%, and nectar volume per flower averages 16.9 μl. The plant populations bloom for one month to year-round, and their flowering approaches the steady-state pattern. Four flower subsets may be defined within the assemblage, each subset related to the bill size and foraging habits of the most frequent bird visitor. Of the six species of hummingbirds recorded at the study site, four are common and largely resident. The four hummingbirds differ in bill size, body mass, and favoured foraging sites, attributes which reflect their favoured flower subsets. One hermit and one trochiline hummingbird share most of the flower species they use, these two birds being the major pollinators within the flower assemblage. This montane forest community may be viewed as medium-rich in ornithophilous flower species and poor in hummingbird species.     

Testosterone levels in feces predict risk‐sensitive foraging in hummingbirds

Risk taking decisions related to the unpredictability of resource availability (risk‐sensitive foraging theory) have typically been explained by behavioral ecology and psychology approaches. However, little attention has been given to the physiological condition of animals as a factor that can influence the direction of foraging preferences. We evaluated the role of steroid hormones testosterone (T) and corticosterone (CORT) on the foraging preferences expressed by white‐eared hummingbirds Hylocharis leucotis in a risk‐sensitivity experiment. We recorded choices made by male individuals to floral arrays with constant and variable rewards (sugar concentration), and associated these with steroid hormone levels quantified at the start of the experiments. We found that males with higher T levels behave as risk‐prone foragers as they perform more visits to flower arrays with variable rewards. Interestingly, CORT levels were similar regardless whether individuals visited both types of array. According to our results, T seems to influence the foraging preferences of male hummingbirds. Individuals with higher levels of this hormone, made more rapid, frequent visits to flowers with variable rewards, and behave consistently as risk‐prone foragers, compared to males with low T levels. These are exciting avenues for future work, particularly considering recent evidence that individuals may exhibit behavioral differences, denoting an apparent personality, which may be associated with phisiological condition influencing how they respond behaviorally to environmental variation.     

Foraging preferences: Response to reward skew

Recent theoretical and empirical work on foraging behaviour suggests that animals may respond to both the means and variances in benefits associated with available resources. We attempt to extend this analysis by asking if reward skew (third moment about the mean) might influence preference when two options have equal means and equal variances. We examine how minimizing the probability of starvation might induce response to skew. In the Appendix we develop an expected ‘fitness’ model which follows from economic theory and indicates more general conditions concerning responses to skew. We also report experiments involving foraging white-crowned sparrows (Zonotrichia leucophrys). Under conditions where positive skew should be favoured, the birds' behaviour supports the prediction. However, their response to skew is not as strong as responses to variance noted in the same individuals.     

Context-dependent foraging choices in risk-sensitive starlings

Existing models of risk-sensitive foraging assume that animals assign value to different options according to an absolute currency. It follows from this assumption that choices are expected to be both transitive and regular, because the value assigned to an option is independent of its context. I tested these predictions by comparing preferences obtained in binary and trinary contexts. European starlings, Sturnus vulgaris, were trained using an operant paradigm to forage for three options: Constant (C) that provided a fixed number of food pellets; Low variance (L) with a coefficient of variation of 71% in the number of pellets; High variance (H) with a coefficient of variation of 106%. The preferences of the birds were tested in three binary choice treatments (CL, CH, LH) and one trinary choice treatment (CLH). Overall, there was no evidence for violations of either transitivity or regularity. However, overall, a bird's relative preference for its most preferred option over its second most preferred option was significantly greater in the trinary treatment than in the comparable binary treatment. This effect of context on choice is compatible with starlings' use of comparative instead of absolute currencies in decision making. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

SELECTIVE PREDATION BY TWO SWIFTS AND A SWALLOW IN CENTRAL AMERICA

Samples of flying insects were made at three sites and during dry and wet seasons in Costa Rica and at two sites during the wet season in Panama. Compared with these were the diets of three aerially foraging birds: the Band-rumped Swift Chaetura spinicauda and Rough-winged Swallow Stelgidopteryx ruficollis in Costa Rica and the Short-tailed Swift Chaetura brachyura in Panama. Among insects flying, Diptera accounted for between 52 and 75%, with smaller numbers of Hymenoptera (mean = 15%) and Coleoptera (10%). In bird stomach contents, however, Hymenoptera predominated (59–81%), Coleoptera were less common (mean = 18.5%) and Diptera least common (3.4%) of the three orders. The discrepancy between diets and samples of flying insects could not be accounted for solely on the basis of differential digestion. The composition of the swift and swallow diets was similar; there were no significant seasonal differences in the taxonomic composition of the diet of Chaetura spinicauda. Size distributions of beetle prey fitted log-normal distributions in every case. Mean size of prey was significantly larger than that of insects available in all comparisons. Although there was no seasonal difference in the mean size of prey of Chaetura spinicauda, the variance in prey size, used as a measure of niche width, was significantly larger in the wet than in the dry season. Although insect densities may be higher, foraging time is reduced and the wet season is likely the worse season. This result therefore supports Emlen's hypothesis of narrower niches under optimal conditions and broader niches under suboptimal conditions. For increasing altitude above ground published data show a decrease in the density of insects flying, a decrease in the proportions of taxa favoured by the birds, and a significant decrease in the mean size of beetles. None of these trends can explain the differences between the birds' diets and the samples of flying insects. Preference of prey by these birds is concluded not to be a function of prey size alone, but to involve interaction among prey size, ease of capture, and local density. Because of different flight abilities, insect taxa will differ in ease of capture and, consequently, in mean size preferred by the predator, as well as proportions in the diet. This factor favours Hymenoptera and Coleoptera over Diptera as prey. Local concentrations of winged ants, bees and social wasps, and fig wasps make these prey differentially easy to exploit and explain the preference of Hymenoptera over Coleoptera. Large species of swifts are hypothesized to be the evolutionary consequence of the tendency of aerially foraging birds to exploit local concentrations of single prey species.     

The foraging decisions of a central place foraging seabird in response to fluctuations in local prey conditions

During reproduction, seabirds need to balance the demands of self- and offspring-provisioning within the constraints imposed by central place foraging. To assess behavioral adjustments and tolerances to these constraints, we studied the feeding tactics and reproductive success of common murres (also known as common guillemots) Uria aalge , at their largest and most offshore colony (Funk Island) where parents travel long distances to deliver a single capelin Mallotus villosus to their chicks. We assessed changes in the distance murres traveled from the colony, their proximate foraging locations and prey size choice during two successive years in which capelin exhibited an order of magnitude decrease in density and a shift from aggregated (2004) to dispersed (2005) distributions. When capelin availability was low (2005), parental murres increased their maximum foraging distances by 35% (60 to 81 km) and delivered significantly larger capelin to chicks, as predicted by central place foraging theory. Murres preferred large (>140 mm) relative to small capelin (100–140 mm) in both years, but unexpectedly this preference increased as the relative density of large capelin decreased. We conclude that single prey-loading murres target larger capelin during long foraging trips as parents are 'forced' to select the best prey for their offspring. Low fledgling masses suggest also that increased foraging time when capelin is scarce may come at a cost to the chicks (i.e. fewer meals per day). Murres at this colony may be functioning near physiological limits above which further or sustained adjustments in foraging effort could compromise the life-time reproductive success of this long-lived seabird.     

Bias in the heritability of preference and its potential impact on the evolution of mate choice

The evolution of mate choice is a function of the heritability of preference. Estimation in the laboratory is typically made by presenting a female with a limited number of males. We show that such an approach produces a downwardly biased estimate, which we term the heritability of choice. When preference is treated as a threshold trait then less biased estimates are obtained particularly for preferences based on the relative value of the preferred trait. Because females in the wild typically survey on average less than five males we argue that the heritability of choice may be more meaningful than the heritability of preference. The restricted number of males surveyed can lead to a reduction in the phenotypic variance of the preferred trait in the group of males selected by the females if the phenotypic variance in preference is equal to or less than the phenotypic variance in the referred trait. If the phenotypic variance in preference exceeds that of the preferred trait then the opposite occurs. A second effect of the restricted number of males sampled is that females are likely to mate initially with males that are not the most preferred. The failure to find the most preferred male may account for the common observation of multiple matings and extra-pair copulations. We suggest that current explanations for polyandry need to take this failure into account.     

Semi‐intensive shrimp farms as experimental arenas for the study of predation risk from falcons to shorebirds

Varying environmental conditions and energetic demands can affect habitat use by predators and their prey. Anthropogenic habitats provide an opportunity to document both predation events and foraging activity by prey and therefore enable an empirical evaluation of how prey cope with trade‐offs between starvation and predation risk in environments of variable foraging opportunities and predation danger. Here, we use seven years of observational data of peregrine falcons Falco peregrinus and shorebirds at a semi‐intensive shrimp farm to determine how starvation and predation risk vary for shorebirds under a predictable variation in foraging opportunities. Attack rate (mean 0.1 attacks/hr, equating 1 attack every ten hours) was positively associated with the total foraging area available for shorebirds at the shrimp farm throughout the harvesting period, with tidal amplitude at the adjacent mudflat having a strong nonlinear (quadratic) effect. Hunt success (mean 14%) was higher during low tides and declined as the target flocks became larger. Finally, individual shorebird vigilance behaviors were more frequent when birds foraged in smaller flocks at ponds with poorer conditions. Our results provide empirical evidence of a risk threshold modulated by tidal conditions at the adjacent wetlands, where shorebirds trade‐off risk and rewards to decide to avoid or forage at the shrimp farm (a potentially dangerous habitat) depending on their need to meet daily energy requirements. We propose that semi‐intensive shrimp farms serve as ideal “arenas” for studying predator–prey dynamics of shorebirds and falcons, because harvest operations and regular tidal cycles create a mosaic of foraging patches with predictable food supply. In addition, the relatively low hunt success suggests that indirect effects associated with enhanced starvation risk are important in shorebird life‐history decisions.     

Foraging information affects the nature of competitive interactions

If animals can learn environmental cues, how might their use of information affect competitive interactions between species? I used shared‐preference isoleg theory to generate four different predictions regarding possible ways information could affect density‐dependent habitat selection. To test these predictions, I conducted field experiments on the foraging behavior of three coexisting species of hummingbirds. I studied black‐chinned (Archilochus alexandri), blue‐throated (Lampornis clemenciae), and magnificent (Eugenes fulgens) hummingbirds at the Southwestern Research Station in the Chiricahua Mts. of Southeastern Arizona, USA. Blue‐throated hummingbirds behaviorally dominate the other two species. I found that birds foraging with complete information (via learned color cues) avoid some of the negative effects from competition. Birds that foraged with complete information remained highly selective on rich feeders even with high competitor densities. When birds suffered an information deficit, however, the two subordinate species shifted their foraging preferences. The dominant species did not shift preference. Each subordinate's shift reflects its unique place in this competitively structured guild. With high competitor densities, black‐chinneds shifted from selective to opportunistic foraging when they also suffered from an information deficit. Thus, an information deficit caused black‐chinneds to cross over their first isoleg. Rather than reduce foraging efficiency, the magnificents shortened their foraging time. I use the magnificents’ response to competition and an information deficit to explore the possibility that this guild of hummingbirds is centrifugally organized. My results require us to consider the information‐gathering (e.g., learning) abilities of individual decision‐makers when we evaluate density‐dependent habitat selection. Differences in the level of usable information can alter the way coexisting species respond to competitive interactions that potentially structure communities.     

The foraging behaviour of Australian honeyeaters: a review and some comparisons with hummingbirds

The foraging behaviour of Australian honeyeaters is reviewed in terms of diet, foraging selectivity, foraging flight mode, quality and quantity of nectar encountered per flower, flower densities encountered and effect of predation. At the same time comparisons are made between honeyeaters and hummingbirds. These two groups of birds are superficially similar. Both feed on nectar and insects. Both tend to have long curved bills and tongues adapted for removal of nectar from flowers. Both tend to feed at long, red flowers. However, on close inspection, honeyeaters and hummingbirds are quite dissimilar. For example, many honeyeaters include fruit in their diets. Hummingbirds almost never eat fruit. Honeyeaters appear to be considerably less nectarivorous and more insectivorous than hummingbirds. Honeyeaters are, for the most part, larger than hummingbirds and they usually perch while feeding whereas hummingbirds usually hover. Honeyeaters but not hummingbirds often flock while feeding. Predation appears to be considerably more important for honeyeaters than for hummingbirds. Territorial defense of flowers seems common in hummingbirds but uncommon in honeyeaters. These differences are discussed in detail and explanations are offered for them wherever possible.     

Relative efficiency of preferred and nonpreferred patterns of lateralized foraging in the gentle lemur (Hapalemur griseus)

Lateralized patterns of hand use in species-typical bamboo shoot foraging were evaluated for efficiency in five female and six male gentle lemurs (Hapalemur griseus sp.). Efficiency was defined as amount of time required to complete a foraging response sequence. The foraging pattern consisted of four component movements: PULL-IN, COUNTERFORCE, TURN, and FEED-IN. These component movements had been shown in a previous study to incorporate lateral hand biases that formed subsets of patterns that were characteristic for each gentle lemur. The duration of each foraging sequence was measured from the beginning of the first component to the initiation of the terminal component. Frequency of use scores were employed to divide the pattern sequences of each subject into preferred and nonpreferred categories. A within-subjects comparison of the mean durations of preferred patterns (M = 2.56 sec) with those of non- preferred patterns (M = 3.02 sec) found that preferred patterns were executed more rapidly, t(10) = 3.36, P = .007. A multiple regression showed that order of pattern preference accounted for 89% of the variance in mean duration of response time (R² = .89, P = .056). Thus, the use of preferred lateralized hand patterns resulted in more rapid bamboo shoot harvesting. Speed in foraging may be regarded as an adaptive strategy for a species that subsists on a high bulk, low nutrient density food such as bamboo. © 1995 Wiley-Liss, Inc.     

A diffusion‐based approach to stochastic individual growth and energy budget,with consequences to life‐history optimization and population dynamics

Using diffusion processes, I model stochastic individual growth, given exogenous hazards and starvation risk. By maximizing survival to final size, optimal life histories (e.g. switching size for habitat/dietary shift) are determined by two ratios: mean growth rate over growth variance (diffusion coefficient) and mortality rate over mean growth rate; all are size dependent. For example, switching size decreases with either ratio, if both are positive. I provide examples and compare with previous work on risk‐sensitive foraging and the energy–predation trade‐off. I then decompose individual size into reversibly and irreversibly growing components, e.g. reserves and structure. I provide a general expression for optimal structural growth, when reserves grow stochastically. I conclude that increased growth variance of reserves delays structural growth (raises threshold size for its commencement) but may eventually lead to larger structures. The effect depends on whether the structural trait is related to foraging or defence. Implications for population dynamics are discussed.     

Comparison of Diuraphis noxia resistance in wheat isolines and plant introduction lines

A bioassay to examine the foraging behaviour of the aphidophagous hoverfly, Episyrphus balteatus was used in which various stimuli purported to influence flower choice were tested using model flowers. In choice experiments, E. balteatus exhibited enhanced behavioural responses to the colour yellow, as previously seen in Eristalis by several workers. Previous suggestions that the generalist forager E. balteatus has no preference for colour are re-examined in the light of this new evidence. The other advertisement cue tested, size, was also demonstrated to play an important role in determining foraging behaviour, with the smaller artificial flowers seemingly preferred. Of the rewards tested, E. balteatus showed preference for the greatest nectar concentrations, whilst increasing pollen did not affect behaviour. These bioassays provided an opportunity to isolate the individual components of decision-making by E. balteatus during foraging. The separation of sexes and ages in these experiments permitted analysis of inter-sexual and inter-generation differences in behaviour, a factor apparently not investigated in previous work.     

Risk-Sensitive Foraging, Fitness, and Life Histories: Where Does Reproduction Fit into the Big Picture?

Risk-sensitive foraging may occur whenever feeding success hasnon-linear effects on fitness. Models of sensitivity to variationin amount of food obtained have concentrated on foraging inorder to survive even though feeding during growth, migration,or reproduction can have strong and non-linear effects on fitness.I illustrate how risk-sensitive foraging for reproduction coulddiffer from risk-sensitive foraging for survival using two simplemodels. Each model assumes that organisms must accumulate somethreshold amount of resources before they can reproduce. Inthe first model, additional resources above the threshold leadto increased reproductive success. Here variance in feedingsuccess can be advantageous even when the mean gain would alloworganisms to reproduce. In the second model, early breedingis superior to late breeding because recruitment rate declinesover time. Here a symmetrical distribution in foraging ratesresults in a skewed distribution of breeding times. Despitethis, variance in feeding success may be advantageous even whenthe seasonal decline in recruitment is linear. The advantageof risk is much larger if foragers can switch freely betweenfeeding options. These two models are united by "the bankruptcyeffect" because all failures to reproduce carry the same consequences.These simple models suggest that risk-prone foraging may belikely when factors other than survival are important for fitness.Finally, I advocate using a mix of modelling techniques, notethe implicitly risk-sensitive nature of stochastic dynamic programming,and speculate on potential connections between risk-sensitiveforaging and theories of life histories in variable environments.INTRODUCTION visit restaurants that provided twice     

Hummingbird incubation: Female attentiveness and egg temperature

Summary Incubating hummingbirds adjust nest attentiveness patterns in different habitats to permit both regulation of egg temperatures for embryonic development and foraging of the adult for positive energy balance. Anna's (Calype anna) and Black-chinned (Archilochus alexandri) Hummingbirds nesting in California chaparral left the nest six to nine times during each daylight hour. Eggs usually cooled only 3–6°C during absences but sometimes cooled up to 20°C during long absences. One Anna's Hummingbird became torpid for 4.5 h at night following a rainy day; the eggs survived cooling to 11°C and hatched two days later. A Costa's Hummingbird (Calypte costae) nesting in the California desert also left the nest several times per hour in early morning and late afternoon, but shaded the eggs almost continuously during the middle of the day. A Purple-crowned Fairy (Heliothryx barroti) in the warm lowland tropics of Panama left her eggs unattended for a few relatively long periods each day rather than many short periods as do temperate zone species. This pattern is typical of other tropical hummingbirds as well probably because equable ambient temperatures mean thermoregulation of eggs is not as critical a problem as it is in other habitats. In the temperate zone, hummingbirds exhibit behavioral adaptations (timing of reproduction, segmented foraging pattern) and physiological adaptations (torpor and hypothermia) for successful incubation. Embryonic development is successful even when egg temperatures fluctuate widely.