Resource use,energetic profitability,and behavioral decisions in migrant rufous hummingbirds

Summary Migrant Rufous Hummingbirds (Selasphorus rufus) defend nectar resources at stopover sites while replenishing fat reserves needed for migratory flights. During late summer in the Sandia Mountains, central New Mexico, they defend the wasp- and bee-pollinated Scrophularia montana from other hummingbirds. Both hummingbirds and hymenopterans exploit Scrophularia nectar during the early part of its flowering period. As summer colony growth increases the densities of the eusocial hymenopterans by 100–150%, their exploitation of Scrophularia nectar lowers its mean standing crop in flowers by 200–300%. Sometime during the summer, Rufous Hummingbirds abandon and do not further use this resource for the remaining 3–4 weeks of its flowering period. The abandonment always occurs when the mean standing crop of nectar is approximately 0.2–0.3 L/ flower. This paper describes a model of Rufous Hummingbird energetics, that shows abandonment occurred 1–3 days after they passed the threshold at which the resource could have provided their minimum daily energy requirements. I suggest that constraints imposed by a highly competitive social environment severely reduced the options available to the hummingbirds, and caused them to continue to defend a resource that could no longer meet their energetic requirements.     

Energy and water balance in the lesser double-collared sunbird (Nectarinia chalybea) feeding on different nectar concentrations

The water balance of nectarivores is tightly linked to their energy balance. When nectar is dilute, consumption of a large water excess is inevitable. We investigated energy and water balance in lesser double-collared sunbirds, Nectarinia chalybea (8 g), kept at 20 °C and fed different nectar concentrations (0.4, 0.8 M sucrose or 1.2 M sucrose). The mass of sucrose consumed, body mass, day-time mass gain and night-time mass loss were the same irrespective of diet, the birds compensating energetically for changes in sucrose concentration by drinking greater volumes of the more dilute solutions. Sunbirds consumed between 0.5 times and 1.8 times their body mass in preformed water per day, depending on sucrose concentration, and excreted around 75% of the water. The difference between water gain (preformed and metabolic water) and excreted water is assumed to equal evaporative water loss, and was similar on 1.2 M and 0.8 M sucrose, but was higher on a diet of 0.4 M sucrose. The osmolalities and K⁺ and Na⁺ concentrations of the excreted fluid were extremely low, so that sunbird urine resembled that of hummingbirds and freshwater vertebrates rather than that of typical terrestrial vertebrates. N. chalybea is able to maintain energy and water balance over a range of nectar concentrations by adjusting the volume of solution consumed and by excreting copious, dilute fluid. Accepted: 2 January 1999     

Flower choice by honey bees (Apis mellifera L.): sex-phase of flowers and preferences among nectar and pollen foragers

Bees foraging for nectar should choose different inflorescences from those foraging for both pollen and nectar, if inflorescences consist of differing proportions of male and female flowers, particularly if the sex phases of the flowers differ in nectar content as well as the occurrence of pollen. This study tested this prediction using worker honey bees (Apis mellifera L.) foraging on inflorescences of Lavandula stoechas. Female flowers contained about twice the volume of nectar of male flowers. As one would predict, bees foraging for nectar only chose inflorescences with disproportionately more female flowers: time spent on the inflorescence was correlated with the number of female flowers, but not with the number of male flowers. Inflorescence size was inversely correlated with the number of female flowers, and could be used as a morphological cue by these bees. Also as predicted, workers foraging for both pollen and nectar chose inflorescences with relatively greater numbers of both male and female flowers: time spent on these inflorescences was correlated with the number of male flowers, but not with the number of females flowers. A morphological cue inversely associated with such inflorescences is the size of the bract display. Choice of flowers within inflorescences was also influenced predictably, but preferences appeared to be based upon corolla size rather than directly on sex phase.     

Dynamics of forager arrivals and nectar renewal in flowers of Anchusa strigosa

Summary Long-tongued Anthophora spp. bees collecting nectar from flowers of Anchusa strigosa (Boraginaceae) exhibit systematic foraging. Successive forager arrivals at individual flowers are not independent, and the time elapsed between successive arrivals at a particular flower is distributed more uniformly than be expected on the basis of a random arrival process. Distributions of inter-arrival time for individual flowers show standard deviation/mean ratios of 0.44–0.79, a range which is consistent with results obtained for two other plant-pollinator systems. The rate at which nectar is renewed between successive forager arrivals is independent of the amount of nectar in the flower, and the renewal process is strongly linear. Practical and theoretical implications of these results are discussed.     

Generous-like flowers: nectar production in two epiphytic bromeliads and a meta-analysis of removal effects

Animal-pollinated angiosperm plants that respond positively to nectar removal by replenishment invest energy that can entail a reproductive cost. We investigated whether or not nectar removal stimulates replenishment in two hummingbird-pollinated bromeliad species. Nectar replenishment rates were also assessed by removing nectar from manually pollinated flowers because pollination events might be used as signals to save energy by preventing allocation to post-pollination nectar production. Then we synthesized the current understanding of nectar removal effects by reviewing existing published studies with a meta-analysis. The magnitude and significance of estimated nectar removal effects and factors associated with variation in size and direction of nectar removal effects were elucidated with the meta-analysis. We found that both Tillandsia species strongly respond to repeated nectar removal by producing >3 times additional nectar. Nectar secretion patterns were not altered by pollination (stigmatic pollen deposition) and we found no evidence of nectar reabsorption. Although the effect size varied widely across systems and/or environmental conditions, the meta-analysis showed that nectar removal had overall a positive effect on nectar replenishment (mainly among species inhabiting wet tropical habitats such as Tillandsia), and a negative effect on the secretion of additional sugar, suggesting that those plants are resource limited and conservative in the secretion of additional sugar.Electronic Supplementary Material Supplementary material is available for this article at     

Energy intake and utilisation by nursing bearded seal (Erignathus barbatus) pups from Svalbard,Norway

In this study we measure energy intake via milk in nursing bearded seal (Erignathus barbatus) pups and determine how this energy is allocated into metabolism and storage of new tissues. This was accomplished using longitudinal mass gain records and the doubly labelled water technique on nursing pups in combination with cross-sectional data on changes in milk composition from bearded seal mothers. The pups (n=3) were all less than a week old at the start of the experiments. Pups gained 3.3±0.4 kg·day^-1 of which 50% was fat, 14% protein and 36% water. Average daily water influx for the pups was 69.5±9.0 ml · kg^-1· day^-1. Average CO₂ production during the study period was 0.99±0.10 ml·g^-1·h^-1, which corresponds to a field metabolic rate of 642±67 kJ·kg^-1· day^-1, or 6.0±0.5 times the predicted basal metabolic rate according to Kleiber (1975). The pups drank an average of 7.6±0.5 kg of milk daily. This corresponds to a daily energy intake of 154±8 MJ, 47±14% of which was stored as new body tissue. Despite this high energy intake bearded seal pups do not get as fat as do other nursing phocids. This is in part due to their larger body size but also due to their very active aquatic lifestyle and the lower and more consistent fat content of the milk compared to other phocid species. Bearded seal mothers forage during lactation and may also be involved in teaching their pups to feed independently. All these data suggest that the lactation strategy of bearded seals differs from the phocid norm.     

Energy allocation rules inDaphnia magna: clonal and age differences in the effects of food limitation

Summary The allocation of energy to carapace formation, respiration, growth, and reproduction were examined in two parthenogenetic clones ofDaphnia magna (Cladocera) cultured at two levels of food (Chlorella) concentration. Clonal differences in energy allocation were more apparent at high ration (1.5 g C mL^-1) than at low ration (0.3 g C mL^-1). These differences included respiratory and molting costs, and the timing of energy allocation to growth and reproduction. A comparison of active vs. anesthetized animals revealed that the interclonal difference in respiration rate was the result of a difference in activity level. In both clones mass-specific rates of respiration, growth, and brood production all decreased at low vs. high ration levels, whereas mass-specific molt-loss rate increased. Lowered food concentration decreased the relative allocation of energy to growth and reproduction, but increased allocation to maintenance (respiration and carapace formation). These allocation responses to food limitation indicated that for both clones the highest energy priority was carapace formation. However, the relative priority of respiration, growth and reproduction varied with age and clone. In juveniles (instars 1–4) the priority ranking of growth was essentially equal to that of respiration, whereas respiration always had higher priority in adults (instars 5–9). All three possibilities for the relative ranking of growth and reproduction (i.e., growth>reproduction, growth=reproduction, and reproduction>growth), as specified by different models in the literature, were observed depending on age and clone. The energy allocation rules were also shown to vary between other daphniid species. Furthermore, metabolic responses to chronic food limitation may be different from responses to acute food deprivation. In this study, one clone showed a greater decrease in respiration rate as a result of lifetime food limitation than did the other, but the opposite was true when these clones were exposed to 48 h of starvation. These differences in allocation rules and in acute vs. chronic responses may have to be considered when using physiological data to modelDaphnia populations.     

Nectar concentration preferences and sugar intake in the white-bellied sunbird, <Emphasis Type="Italic">Cinnyris talatala</Emphasis> (Nectariniidae)

Plant nectar is a simple food consumed by many different animals. Preferences regarding its components, especially sugars, have been studied for many species, but the preferences of nectar-feeding birds for different sugar concentrations are less well known than their sugar type preferences. The concentration preferences of white-bellied sunbirds, Cinnyris talatala, were examined using paired solutions of either sucrose or equicaloric 1:1 mixtures of glucose and fructose. Preferences were tested over a broad concentration range of 0.25–2.5 M sucrose equivalents (using 0.25 or 0.5 M differences between pairs). On both sucrose and hexose diets, the higher concentration was preferred up to 1 M, but there were no significant preferences above this concentration, except that birds preferred 1.5 to 2 M sucrose. As with other nectar-feeding vertebrates, the laboratory preferences of sunbirds do not explain the low concentration of their natural nectars. We recorded apparent excess sugar consumption during 6 h preference tests involving concentrated hexose diets; this could be due to digestive constraints or viscosity differences between sucrose and hexose solutions.     

Nectary structure and nectar characteristics in someBignoniaceae

The nectary structure and chemical nectar composition of 15 species belonging to 12 genera ofBignoniaceae are analyzed. All taxa bear a conspicuous nuptial nectary surrounding the ovary base. The secretory tissue is mostly supplied by phloem branches. The stomata are located in the middle and upper part of the nectary epidermis with an homogeneous distribution. The nuptial nectary is proportionally large in relation to the ovary (15–30%), disregarding the nectary volume. Most species have extranuptial nectaries in both inner and outer surfaces of the calyx. Both kinds of nectaries lack a vascular tissue that straightly supplies them. Nuptial nectar concentration (wt/wt) ranges from 19 to 68%. Sugars and amino acids are found in all species. Half of the species have hexose predominant nectars, the remaining sucrose predominant. Phenols are detected in only three species, whereas reducing acids exclusively inTecoma stans. Alkaloids and lipids were never detected. Extranuptial nectar chemical composition is analyzed in two species:Dolichandra cynanchoides andPodranea ricasoliana. Bees constitute the main flower visitors of the species studied whereas hummingbirds were seen visiting three species. A correlation analysis is performed with the data obtained. There are a few significant correlations which indicate a parallel increase of three parameters: the longer the flower length, the more voluminous the nectary and the higher stomata number, independently of the floral biotype. Phenograms are obtained using 24 floral characters including nectary and nectar data. The clusters obtained do not reflect taxonomic relationships but are useful in the understanding of animal-plant interactions when the flower biotype is considered.This paper is based on a chapter of a doctoral thesis presented at the University of Córdoba (Argentina).     

Absence of nitrogen fixation (acetylene reduction) by procaryotes in nectar of Banksias

Summary An earlier hypothesis that blue-green algae in the nectar ofBanksia telmatiaea contribute to the nitrogen economy of the host by fixing N₂ was tested. Field and laboratory experiments failed to demonstrate C₂H₄ production in C₂H₂-treated containers over extended periods. Soil N was not higher at the end of the flowering season and plants in which flower heads were removed prior to nectar production did not contain less N than the controls.     

Variation in nectar volume and composition of Impatiens capensis at the individual,plant, and population levels

Although the volume and chemical composition of nectars are known to vary among plant species and to affect pollinator response to plants, relatively little is known of the variation in volume, and sugar and amino acid composition within species. We collected nectar from Impatiens capensis in a nested design: three flowers from each of three plants from each of three populations. This design enabled us to quantify variation within individual plants, among plants within populations, and among populations. Using high performance liquid chromatography, we analyzed the sugar and amino composition of the 27 flowers. Analysis of variance showed that none of the parameters (volume, concentrations of three sugars and 24 amino compounds) varied within individuals. Variation in nectar volume was not significant among plants but was nearly significant among populations. Of the three sugars detected (sucrose, glucose, and fructose), the only significant variation was that of sucrose among populations. Concentrations of 12 amino compounds varied significantly at the plant level while 7 amino compounds varied among populations. The results indicate that: (1) pooling of nectar samples from flowers of individual plants can be an acceptable methodology for those seeking to understand within species variation; (2) amino compounds appear to vary more than either volumes or sugar concentrations; (3) future studies should assess how much of the observed variation is due to genetic versus environmental differences; (4) additional studies should examine the geographic variation in nectar parameters and pollinators of I. capensis in order to assess the role different pollinators play in shaping nectar composition.     

Genetics of flower size and nectar volume in Petunia pollination syndromes

The two related Petunia species, P. axillaris and P. integrifolia, are sympatric at various locations in South America but do not hybridise. Divergent pollinator preferences are believed to be in part responsible for their reproductive isolation. The volume of nectar produced and several components of flower morphology might contribute to pollinator-dependant reproductive isolation. In this study, we aimed to identify the genetic changes underlying the quantitative differences observed between these two Petunia species in flower size and nectar volume. We mapped quantitative trait loci (QTL) responsible for the different phenotypes of P. axillaris and P. integrifolia in an inter-specific backcross population. QTL of small to moderate effect control the differences in flower size and volume of nectar. In addition, we observed strong suppression of meiotic recombination in Petunia, even between closely related species, which precluded a fine resolution of QTL mapping. Thus, our data suggest that flower size and nectar volume are highly polygenic. They are likely to have evolved gradually through pollinator-mediated adaptation or reinforcement, and are not likely to have been primary factors in early steps of pollinator isolation of P. axillaris and P. integrifolia.     

Adult feeding by holometabolous insects: pollen and nectar as complementary nutrient sources for Rhingia campestris (Diptera: Syrphidae)

Summary Examination of the gut contents of adult Rhingia campestris reveals that there are considerable differences in the relative amounts of pollen and nectar ingested by individual insects. Pollen, a rich protein source, is required most by females, particularly during those stages of ovarial development when yolk deposition occurs. Nectar, which is mainly carbohydrate, is required in large amounts by males, and is also required by females before and after oogenesis. These results are discussed in relation to the likely nutritional needs of the flies, and it is suggested that similar feeding patterns may be found in a variety of other flower visiting insects. Finally, reference is made to the problems of making ecological assumptions about feeding behaviour on the basis of the morphological attributes of the mouthparts.     

Variability in nectar production and standing crop, and their relation to pollinator visits in a Mediterranean shrub

Nectar standing crops in flowers within an individual plant are often highly variable. This variability may be a by-product of the foraging activity of insect pollinators. Alternatively, plants may be selected to produce highly variable rewards to reduce consecutive visitation by risk-averse pollinators, thus diminishing within-plant pollen transfer. This study evaluated the roles of pollinator control vs. plant control over nectar variability in the bee-pollinated shrub Rosmarinus officinalis L. (Lamiaceae). We sampled nectar production, standing crop and pollinator visits in three shrubs of one population over 17 days during one blooming season. Nectar production rates were highly variable (CV = 1.48), and increased after rainy days. Nectar standing crops were even more variable (CV = 2.16), decreased with increasing temperatures, and increased with time since the last rain. Pollinator visit rates decreased with variability in nectar standing crops, increased with flower number per shrub, and were unaffected by variability in nectar production rates. Repeated sampling of marked flowers revealed no correlation between their nectar standing crops and production rates. These findings support the role of reward variance in reducing pollinator visits, but suggest that plants are not in complete control of this variability. Rather, plant-generated variability can be modified by intensive foraging activity of pollinators. Such pollinator control over nectar variability is likely to reduce the selective advantage of plant-generated reward variation. Handling Editor: Neal Williams.     

Patch selection by red deer in relation to energy and protein intake: a re-evaluation of Langvatn and Hanley's (1993) results

Langvatn and Hanley (1993) recently reported that patch use by red deer (Cervus elaphus) was more strongly correlated with short term rates of intake of digestible protein than dry matter. Such short term measures overlook effects of gut filling, which may constrain intake by ruminants over longer time scales (i.e., daily rates of gain). We reanalyzed Langvatn and Hanley's data using an energy intake model incorporating such a processing constraint, to determine whether their conclusions are robust. We found that the use of patches by red deer was just as strongly correlated with an estimate of the daily rate of intake of digestible energy as one of digestible protein during four out of seven trials, but slightly lower in three out of seven trials. In all cases, daily intake of digestible energy was a much better predictor of patch preference by red deer than was the intake of dry matter. Our reanalysis suggests that the daily intake of energy was highly correlated with that of protein in these trials, as may often be the case for herbivores feeding on graminoids. Hence the observed pattern of patch use by red deer could simultaneously enhance rates of both protein and energy intake.     

Nectar by day and night:Siphocampylus sulfureus (Lobeliaceae) pollinated by hummingbirds and bats

Most species of the neotropical genusSiphocampylus are believed to be bird-pollinated. The pollination biology ofSiphocampylus sulfureus was studied in a montane region in SE Brazil. This species has features intermediate between ornithophilous and chiropterophilous syndromes: it presents a striking combination of yellowish flowers with strong odour (chiropterophilous features), and diurnal anthesis and sucrose-dominated nectar (ornithophilous features). Major pollinators were hummingbirds by day, and a phyllostomid bat by night.Siphocampylus sulfureus may be viewed as a recent derivate from the presumed ornithophilous stock within sect.Macrosiphon, and thus benefits from the activity of both diurnal and nocturnal vertebrate pollinators.Dedicated to DrIlse Silberbauer-Gottsberger and Prof. DrGerhard Gottsberger for their pioneer contribution to floral biology in Brazil.     

Effect of nectar supplementation on male and female components of pollination success in the deceptive orchid Dactylorhiza sambucina

Many orchids lack floral nectar rewards and therefore rely on deception to attract pollinators. To determine the effect that a mutation for nectar production would have on overall pollination success of the deceptive orchid Dactylorhiza sambucina, we recorded pollen deposition and removal in flowers of plants that had either been supplemented with an artificial nectar solution or left unmanipulated as controls. Nectar supplementation resulted in significant increases in the proportion of flowers pollinated, regardless of morph colour and the density of plants supplemented in the population. However, nectar supplementation had a significant positive effect on pollinaria removal only for the yellow morph in one experiment in which a low proportion of plants were supplemented. Thus a mutation for nectar production would have a positive effect on overall pollination success in D. sambucina, particularly the female component. The observed patterns are discussed in relation to other factors, such as cross-pollination and the reallocation of nectar resources for other plant functions, which are traditionally considered to shape the rewardless strategies of orchids.     

Limits to food intake and fiber utilization in the prairie vole,Microtus ochrogaster: effects of food quality and energy need

We fed prairie voles (Microtus ochrogaster) rat chow diluted with variable amounts of -cellulose to determine 1) how much fiber the voles could tolerate in their diet; 2) changes in food intake and digestibility of dry matter and of fiber; 3) the extent to which voles utilized fiber as an energy source; and 4) whether any of these variables differed between groups of animals maintained at 5 or 22°C. Fiber content of the diets ranged from 20 to 84%. Animals held at 5°C maintained body mass through a diet containing 69% fiber, while animals held at 22°C maintained body mass through the 84% fiber diet. Dry matter intake increased with fiber level from 9.3 to 15.0 g·day^-1 for animals at 5°C and from 5.6 to 14.0 g·day^-1 for animals at 22°C; intake on the highest fiber diet eaten by either group was not different. Dry matter digestibility decreased significantly as the fiber in the diets increased, but was not affected by temperature treatments. Digestible dry matter intake for each group remained constant regardless of diet quality, but on each diet digestible dry matter intake for animals at 5°C was significantly higher than that of the animals held at 22°C. Digestibility of the fiber portion of the experimental diets remained constant as food quality decreased, so the percent of daily energy need met by fiber utilization increased with higher food intake. On the lowest quality diet each group tolerated, fiber digestion provided approximately 42 and 68% of the energy needs of voles at 5 and 22°C, respectively.Abbreviations BM body mass - BMR basal metabolic rate - DE digestible energy - DM dry matter - DMD dry matter digestibility - DDMI digestible dry matter intake - MR metabolic rate - NDF neutral detergent fiber (=cell walls) - NDS neutral detergent solubles (=cell solubles) - SEM standard error of mean - T _a ambient temperature     

Development of net energy intake models for drift-feeding juvenile coho salmon and steelhead

We developed models to predict the effect of water velocity on prey capture rates and on optimal foraging velocities of two sympatric juvenile salmonids, coho salmon and steelhead. Mean fish size was ~80 mm, the size of age I+ coho and steelhead during their second summer in Southeast Alaska streams, when size overlap suggests that competition might be strongest. We used experimentally determined prey capture probabilities to estimate the effect of water velocity on gross energy intake rates, and we modeled prey capture costs using experimental data for search and handling times and published models of swimming costs. We used the difference between gross energy intake and prey capture costs to predict velocities at which each species maximized net energy intake rate. Predicted prey capture rates for both species declined from ~75 to 30–40 prey/h with a velocity increase from 0.30 to 0.60 m·s⁻¹. We found little difference between coho and steelhead in predicted optimum foraging velocities (0.29 m·s⁻¹ for coho and 0.30 m·s⁻¹ for steelhead). Although prey capture ability appears to be more important than are prey capture costs in determining optimum foraging velocities, capture costs may be important for models that predict fish growth. Because coho are assumed to pay a greater swimming cost due to a less hydrodynamic body form, we also modeled 10 and 25% increases in hydrodynamic drag to assess the effect of increased prey capture costs. This reduced optimum velocity by 0 and 0.01 m∙s⁻¹, respectively. Habitat segregation among equal-sized coho and steelhead does not appear to be related to the effects of water velocity on their respective foraging abilities.     

Bumblebee visits to Impatiens spp.: pattern and efficiency

Summary Three Japanese species of Impatiens, which secrete nectar continuously in long spurs, were visited by Bombus diversus workers consecutively throughout the day. B. diversus workers showed characteristic patterns of behavior in flower use, flower choice, and patch departure. (1) Bumblebees stayed longer on a flower which had been unvisited for a while than on a flower which had been visited recently. (2) Bumblebees preferred visiting flowers which had been unvisited for a while to visiting those which had been visited recently, and to visiting those which had been unvisited for a long period. (3) Bumblebees had a higher probability of leaving a patch after they had stayed on a flower for a short period than after they had stayed for a longer period. The bumblebees appeared to perceive both remotely and proximately chemical cues deposited by other foraging individuals, which indicated nectar rewards in a flower, and thus obtained a higher nectar intake than the mean amount of nectar left in a flower.