Digestion of nectar and insects by Palestine sunbirds

In nectarivorous birds, specialization for feeding on nectar has led to a simple gut structure with high sugar digestive efficiencies and rapid gut passage rates. These features of the digestive system may make digestion of more complex, protein-rich food sources, such as pollen or insects, less efficient. In this light, we hypothesized that sugar metabolizability in nectarivorous Palestine sunbirds (Nectarinia osea) would be high, whereas nitrogen metabolizability would be lower than typically found for birds. We measured glucose and fructose apparent metabolizabilities (*MCs) and transit times (TTs) in eight Palestine sunbirds offered either a 10% or a 50% mixed sugar diet. *MC for glucose (99.9%+/-0.1%) was significantly greater than for fructose (99.6%+/-0.4%; ANOVA; P<0.001). TT for the 10% sugar diet (26.3+/-10.1 min) was significantly shorter than for the 50% sugar diet (47.0+/-7.8 min). We measured nitrogen true metabolizability (MC) and TT in Palestine sunbirds offered a daily fruit fly intake of either 40 or 200 flies. Nitrogen MC was not significantly different between diets, and average MC for both diets was 58.5%+/-8.5% (n=8). TT was not significantly different when birds ate 10 flies (50.1+/-13.6 min) than when they ate 50 flies (48.5+/-16.5 min). The high sugar *MC and relatively rapid TT of nectar in Palestine sunbirds are similar to those found for other nectarivorous species. Transit times of insect material are longer that those found in small insectivorous species. However, MCs of insect material are lower. Thus, even though sunbirds consume easily digestible soft-bodied insects, they are less efficient at extracting protein than nonnectarivores.     

Lesser double‐collared sunbirds Nectarinia chalybea do not compensate for hatching asynchrony by adjusting egg mass or yolk androgens

Substantial amounts of maternal androgens are found in birds’ eggs and have been shown to benefit offspring development. Within‐clutch patterns of increasing androgen concentrations over the laying sequence are often hypothesized to compensate for the negative effects of hatching asynchrony. However, detrimental effects to offspring fitness of exposure to high yolk androgen levels have also been demonstrated. This suggests that mothers should forego these costs to their offspring when the need for compensation for hatching asynchrony is low or when alternative compensatory strategies, e.g. in terms of increasing egg mass, are available. Here we show that in the south‐temperate lesser double‐collared sunbird Nectarinia chalybea, a species with hatching asynchrony but also with high survival of last‐hatched chicks, mothers do not deposit resources differentially in terms of either yolk androgen concentration or egg mass across the laying sequence. We discuss to what extend this challenges the original explanation of within‐clutch variation in these egg parameters and offer some explanation for their between‐clutch variation which was related to female body mass.     

Dietary intake effects on arginine vasotocin and aldosterone in cloacal fluid of whitebellied sunbirds (Nectarinia talatala)

The main objective of this study was to determine whether or not the renal outputs of the osmoregulatory hormones arginine vasotocin (AVT) and aldosterone (ALDO) reflect the osmotic status of whitebellied sunbirds (Nectarinia talatala). The birds were fed a range of sucrose concentrations (from 0.07 to 2.5 mol/l, with osmolalities of 70 to approximately 5,800 mosM/kg), and adjusted their intakes so that they drank large volumes of dilute diets and small volumes of concentrated diets. Renal fluid outputs were appropriately regulated so that large volumes of cloacal fluid (CF) were voided on the dilute diets and small volumes on the concentrated diets. Accordingly, plasma AVT concentrations increased with increasing sugar concentration; however, AVT outputs in CF did not change in a similar manner, rather they decreased as dietary concentration increased. It was not possible to measure plasma ALDO concentrations in the small sunbirds because of insufficient blood samples available; however, ALDO outputs in CF did vary with the sucrose diets and renal function, being highest on the most concentrated diet. In addition ALDO output in CF fell markedly when sodium was added to the 0.5 mol/l sucrose diet. We conclude that in sunbirds fed increasingly concentrated sucrose solutions, changes in CF outputs of ALDO, but not AVT, appear to reflect the water flux and hydration state of these birds.     

Survival of male scarlet-tufted malachite sunbirds (Nectarinia johnstoni) on Mount Kenya and the influence of ornamentation

A prediction of handicap and signalling theories is that there should be a positive relationship between ornament size and viability. This prediction was tested in scarlet-tufted malachite sunbirds ( Nectarinia johnstoni ). These birds are endemic to the high-altitude zones of East African mountains. They are bright iridescent green with elongated central tail feathers and red pectoral tufts. The elongated tail is believed to be a mate choice cue, and the red tufts play a role in aggressive encounters between males. As adults, they feed almost exclusively on the nectar of Lobelia telekii flowers, which males defend from conspecifics. The long term survival of individually identifiable males colour-marked in 1989 and 1990 was investigated. Territorial adult males had an average annual survival rate of 72%, which is at the top of the range reported for tropical birds. The effects of morphology and territory characteristics on male survival were examined by adding covariates to the survival model. This analysis showed that male N. johnstoni with longer tails had a higher probability of survival than did short-tailed males. There was no apparent relation between tuft size or the number of L. telekii inflorescences on the territory and survival. The results presented here suggest that long-tailed males may be of intrinsically higher quality than are shorter tailed males. If this is true then females may benefit by using this trait as a mate choice cue through increased viability of their offspring.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 125–133.     

Molecular phylogeography reveals island colonization history and diversification of western Indian Ocean sunbirds (Nectarinia: Nectariniidae)

We constructed a phylogenetic hypothesis for western Indian Ocean sunbirds (Nectarinia) and used this to investigate the geographic pattern of their diversification among the islands of the Indian Ocean. A total of 1309 bp of mitochondrial sequence data was collected from the island sunbird taxa of the western Indian Ocean region, combined with sequence data from a selection of continental (African and Asian) sunbirds. Topological and branch length information combined with estimated divergence times are used to present hypotheses for the direction and sequence of colonization events in relation to the geological history of the Indian Ocean region. Indian Ocean sunbirds fall into two well-supported clades, consistent with two independent colonizations from Africa within the last 3.9 million years. The first clade contains island populations representing the species Nectarinia notata, while the second includes Nectarinia souimanga, Nectarinia humbloti, Nectarinia dussumieri, and Nectarinia coquereli. With respect to the latter clade, application of Bremer's [Syst. Biol. 41 (1992) 436] ancestral areas method permits us to posit the Comoros archipelago as the point of initial colonization in the Indian Ocean. The subsequent expansion of the souimanga clade across its Indian Ocean range occurred rapidly, with descendants of this early expansion remaining on the Comoros and granitic Seychelles. The data suggest that a more recent expansion from Anjouan in the Comoros group led to the colonization of Madagascar by sunbirds representing the souimanga clade. In concordance with the very young geological age of the Aldabra group, the sunbirds of this archipelago have diverged little from the Madagascar population; this is attributed to colonization of the Aldabra archipelago in recent times, in one or possibly two or more waves originating from Madagascar. The overall pattern of sunbird radiation across Indian Ocean islands indicates that these birds disperse across ocean barriers with relative ease, but that their subsequent evolutionary success probably depends on a variety of factors including prior island occupation by competing species.     

Seasonal and annual variation in the diet of breeding and non-breeding Wilson's storm-petrels on King George Island, South Shetland Islands

The diet of breeding and non-breeding Wilson's storm-petrels (Oceanites oceanicus) was analysed over four breeding seasons on King George Island, South Shetland Islands, in order to test if there are changes in diet composition within and between seasons. Krill was the most important food item, followed by myctophid fish and amphipods. Breeding birds showed a clear seasonal pattern of diet, with krill decreasing and alternative prey increasing from the incubation to the chick-rearing period. Non-breeders were not found to change their diet composition. The data suggest that Wilson's storm-petrels selectively choose alternative prey to krill in order to meet the nutrient demands of their chicks. In years of low food availability, Wilson's storm-petrels may have a limited option of prey choice.     

Intersexual differences in energy expenditure of Anolis carolinensis lizards during breeding and postbreeding seasons

Although the amount of energy that males and females invest in reproduction is an integral component of theories explaining the evolution of particular mating strategies, few studies have actually determined the amount of energy that each sex allocates to reproduction. We compared how energy is expended by male and female Anolis carolinensis lizards during both the breeding and postbreeding seasons. We used laboratory respirometry to determine resting metabolic rates (RMRs) of inactive, freshly captured lizards and the doubly labeled water technique to determine field metabolic rates (FMRs) of free-ranging lizards. Both RMRs and FMRs were influenced by body mass but not by sex. Season did not influence FMRs; however, RMRs of both sexes increased approximately 40% from the breeding to the postbreeding season. The seasonal increase in RMRs was attributed to a postreproductive increase in feeding rate and specific dynamic action. We used RMRs, FMRs, and thermal profiles of lizards to calculate energy budgets for breeding and postbreeding seasons. Energy budgets partitioned daily field energy (DFE; calculated from FMRs) into daily activity energy (DAE) and daily resting energy (DRE; calculated from RMRs). Energy expended for reproduction was estimated as DAE during the breeding season plus egg production (for females). Despite males having 40% greater body mass, females expended 46% more energy for reproduction than did males (906 and 619 J/d, respectively). Total metabolizable energy (TME=DFE+egg production for females) expended during the breeding season was similar for males and females (1,280 and 1,365 J/d, respectively). Although TME of females decreased 44% from the breeding to the postbreeding season (1,365 vs. 766 J/d), TME of males was similar during both seasons (1,280 vs. 1,245 J/d). There were both seasonal and sexual differences in DRE and DAE. Compared with most lizards from semiarid/desert habitats, A. carolinensis in a temperate habitat expends more total energy during the breeding season, allocates more energy to eggs, and appears to have more total energy available for reproduction.     

New specific equation to estimate resting energy expenditure in severely obese patients

Calculating the estimated resting energy expenditure (REE) in severely obese patients is useful, but there is controversy concerning the effectiveness of available prediction equations (PE) using body weight (BW). We evaluated the efficacy of REE equations against indirect calorimetry (IC) in severely obese subjects and aimed to develop a new equation based on body composition compartments. One hundred and twenty severely obese patients had their REE measured (MREE) by IC and compared to the most commonly used PE (Harris-Benedict (HB), Ireton-Jones, Owen, and Mifflin St. Jeor). In a random sample (n = 60), a new REE equation based on fat-free mass (FFM) was developed and validated. All PE studied failed to estimate REE in severe obesity (low concordance correlation coefficient (CCC) and limits of agreement of nearly 50% of the sample ±10% of MREE). The HB equation using actual BW exhibited good results for all samples when compared to IC (2,117 ± 518 kcal/day by HB vs. 2,139 ± 423 kcal/day by MREE, P > 0.01); these results were blunted when patients were separated by gender (2,771 vs. 2,586 kcal/day, P < 0.001 in males and 1,825 vs. 1,939 kcal/day, P < 0.001 in females). A new resting energy expenditure equation prediction was developed using FFM, Horie-Waitzberg, & Gonzalez, expressed as 560.43 + (5.39 × BW) + (14.14 × FFM). The new resting energy expenditure equation prediction, which uses FFM and BW, demonstrates higher accuracy, precision, CCC, and limits of agreement than the standard PE in patients when compared to MREE (2,129 ± 45 kcal/day vs. 2,139 ± 423 kcal/day, respectively, P = 0.1).The new equation developed to estimate REE, which takes into account both FFM and BW, provides better results than currently available equations.     

Coping with predators and food limitation: testing life history theory for sex–specific larval development

For animals with complex life cycles, recent models of sexual size-dimorphism at maturity assume three key variables to optimise larval life history: activity in the larval stage, development time, and size at maturation. However, model predictions remain largely untested. In the territorial dragonfly Libellula depressa (Odonata) exhibiting a flexible development time we tested for male-biased sexual size-dimorphism and sex differences in larval activity, development time, and growth rate. Based on models we predicted that males achieved their larger size compared to females by a longer development rather than being more active. Results revealed that males took longer to develop and achieved a larger size than females but were not more active. Compared to males, females exhibited a higher growth rate which was not achieved by an activity-mediated increased food intake. We conclude that sexual size-dimorphism in species with a flexible development time is mediated by differences in developmental length but not activity. Furthermore, sexes differ in their plastic responses to food availability and predator presence making it necessary to consider sex-specific differences in testing further life history responses.     

Seasonal variation in energy expenditure and body composition in captive White Storks (Ciconia ciconia)

North Western European populations of White Storks (Ciconia ciconia) appear to have been saved from extinction by settling, i.e. stopping migration. Settled storks exposed to winter conditions must cope with periods of potentially high energy demands that would otherwise be avoided by the migration process. Doubly labeled water (DLW) was therefore used to examine the seasonal variation (summer vs winter) in daily energy expenditure (DEE) and the body composition of adult and immature storks of both sexes. Male White Storks showed a higher DEE over the winter period than in summer compared with females; in particular, immature males exhibited greater energy expenditure in winter than adult males. Thus, the DEE did not significantly differ between summer and winter (except for immature males), reflecting an absence of thermoregulation cost in winter. For both age classes, total body mass increased in winter, which was mainly due to an increase in fat mass. Adult storks were 5% heavier than immature storks. The sexes differed in body mass, with males weighing significantly more than females by 11%. Mean LBM (lean body mass) was 8.5% higher in adults than in immatures, and was 11.5% higher in males compared with females. Between their first and second summers, immatures accumulated a lean body mass to finally reach the same values as adults, indicating a phase of muscle development. The mean fat mass of the storks did not differ between age classes or between sexes. Based on physiological parameters, this study shows that settled White Storks are able to cope with mild winter periods when they are artificially provided with food. In a view to preserve favourable habitats for this species, it is therefore necessary to decide on a plan of action for breeding areas.     

Avian responses to forest fragmentation during the breeding and non-breeding seasons

Forest fragmentation represents a threat to several bird species worldwide. Several factors can change across seasons (e.g. bird perception of the landscape, weather conditions, biotic interactions), which can modify the response of bird populations to forest fragmentation. However, most studies have been conducted only during the breeding season. Here we assessed the relationship between forest fragmentation (patch area and patch isolation) with population abundances of resident species during both the breeding and the non-breeding seasons. Bird population abundances (all species in the community, subsets of forest and habitat generalist species and for individual species) were estimated across a gradient of area-isolation in a semi-arid forest in Cordoba, Argentina. Population abundance of the overall avian community and of the subset of forest species declined with patch area reduction independently of the season. By contrast, the subset of habitat generalist species was not affected by patch area reduction or by the increase in patch isolation, either during the breeding or during the non-breeding season. When the analyses were carried out for individual species, we found four forest species and one habitat generalist species whose responses (the relationship between population abundance and patch area or with isolation) were different between breeding and non-breeding seasons. The negative effects of forest fragmentation were found mainly during the breeding season. Our results suggest that reduction of patch area may lead to a reduction of more than 65% of the population abundance of forest bird species, during both the breeding and the non-breeding season. Therefore, there is an urgent need to conserve large forest patches within the region as irreplaceable elements for the conservation of populations of several species.     

Activity and energy expenditure

The influence of small changes in activity on energy expenditure and hence on energy requirements and energy balance is assessed. Evidence from direct and indirect calorimetry suggests that differences in spontaneous minor activity could readily alter 24-h energy expenditure by as much as 20%. This compares with values in the order of 10% for moderate overfeeding and somewhat less than this during mild cold exposure. Individual variability in 24-h energy expenditure can therefore be accounted for not only by differences in resting metabolism and the thermic responses to energy intake and temperature but also by differences in minor activity. Interactions between activity and environmental factors such as nutrition and temperature can modify the effect of activity on energy balance. Very little is known about mechanisms that could account for differences in spontaneous activity and these need to be the subject of future investigations.     

Tri-axial accelerometry shows differences in energy expenditure and parental effort throughout the breeding season in long-lived raptors

Cutting-edge technologies are extremely useful to develop new workflows in studying ecological data, particularly to understand animal behavior and movement trajectories at the individual level. Although parental care is a well-studied phenomenon, most studies have been focused on direct observational or video recording data, as well as experimental manipulation. Therefore, what happens out of our sight still remains unknown. Using high-frequency GPS/GSM dataloggers and tri-axial accelerometers we monitored 25 Bonelli’s eagles Aquila fasciata during the breeding season to understand parental activities from a broader perspective. We used recursive data, measured as number of visits and residence time, to reveal nest attendance patterns of biparental care with role specialization between sexes. Accelerometry data interpreted as the overall dynamic body acceleration, a proxy of energy expenditure, showed strong differences in parental effort throughout the breeding season and between sexes. Thereby, males increased substantially their energetic requirements, due to the increased workload, while females spent most of the time on the nest. Furthermore, during critical phases of the breeding season, a low percentage of suitable hunting spots in eagles’ territories led them to increase their ranging behavior in order to find food, with important consequences in energy consumption and mortality risk. Our results highlight the crucial role of males in raptor species exhibiting biparental care. Finally, we exemplify how biologging technologies are an adequate and objective method to study parental care in raptors as well as to get deeper insight into breeding ecology of birds in general.     

Seasonal breeding in female mammals

Study of the causation of seasonal breeding can be subdivided into a consideration of external or ecological, and internal or physiological, factors. Of the ecological ones, the role of day–length has been most extensively studied. Subjecting appropriate animals to different photoperiods should give fresh insight into mechanisms of the central nervous system. Alterations in day–length may produce changes in the C.N.S. and pituitary quite quickly, say within 24 to 48 hours. But there is still uncertainty about such basic problems as the relative significance of abrupt and gradual changes in day–length for the alteration of gonadal activity; and the existence of an annual endogenous physiological rhythm which may provoke the gonads into activity at the appropriate time. In addition, for some small rodents physical environmental factors apparently are not always equally efficient in stopping breeding in the winter, and it may be that length of breeding season is also influenced by biotic factors, such as the phase of a population cycle. Susceptibility to the effects of short photoperiods ("winter") may be to some extent genetically controlled. Possibly then genotypes allowing winter breeding may be favoured at some, but not other, phases of population cycles.     

Mutual benefits of associations between breeding and non-breeding White-fronted Geese Anser albifrons

The relationships between yearlings and adult pairs of White-fronted Geese Anser albifrons were studied during pre-nesting, laying and early incubation in the central Canadian Arctic. Prior to nesting, females of lone pairs spent 75–81% of their time feeding, while males spent only 42–47% of time feeding and 46–50% alert. In pairs with one or more associated yearlings, both females and males fed significantly more and spent less time vigilant. Yearlings spent significantly less time (59%) feeding when alone compared with 71–76% when with pairs. Associations between yearlings and paired adults were most frequent before adult females began prospecting for nest sites. No prospecting pairs were associated with yearlings. After the egg-laying period, groups of geese, predominantly yearlings, made distraction flights over humans and terrestrial predators approaching nests, in contrast to the more cryptic behaviour of nesting pairs. The presence of groups of geese associated with some nest sites suggests that continuing parent-offspring relationships may involve assistance with nest defence.     

Seasonal differences in energy expenditure,flight characteristics and spatial utilization of Dalmatian Pelicans Pelecanus crispus in Greece

Animals typically adjust their behaviour to their changing environment throughout the annual cycle, modulating key processes such as the timing of breeding and the onset of migration. Such behavioural changes are commonly manifested in the movements and the energetic balance of individuals in relation to their species‐specific physiological characteristics, habitat attributes and the environmental properties of their distribution ranges. We used GPS and acceleration data collected using transmitters on free‐ranging birds to quantify annual movement patterns and estimate energy expenditure of the Dalmatian Pelican Pelecanus crispus, a large, soaring avian species which performs short‐distance migration and spends its entire annual cycle in mid‐latitudes. To assess the representativeness of our results, the transmitter effect was also tested. We found that daily trends in the overall dynamic body acceleration (ODBA; a proxy for energy expenditure) differed among seasons, with the highest values occurring during spring and the lowest during winter. Long inter‐lake flights were very rare in winter, and the number of flights and ODBA during spring was higher than during summer, suggesting greater motivation to move in spring. Although transmitters may have affected the birds, as none of the tagged birds bred, we found seasonal differences in behaviour and activity level. The observed patterns in differences in activity levels, long‐distance flights and flight characteristics between seasons suggest an annual rhythm of energy expenditure. These findings allow a better understanding of bird phenology, specifically regarding adaptations to wintering in a cold climate by reducing movement‐driven energy expenditure. Finally, the identification of periods with high and low energy expenditure may guide future conservation efforts by adjusting conservation plans in accordance with changing needs during the annual cycle.     

Social control of reproduction in breeding and non-breeding male naked mole-rats (Heterocephalus glaber).

Eight male naked mole-rats, from three colonies were studied in captivity. When non-breeding male naked mole-rats were removed from their colonies and paired with a non-breeding female, or removed and housed singly for 6 weeks before pairing with a female, concentrations of urinary testosterone and plasma luteinizing hormone (LH) increased significantly (P less than 0.05). Concentration of these hormones were highest while the males were singly housed: urinary testosterone (mean +/- s.e.m.) increased from 8.2 +/- 1.3 ng/mg urinary creatinine (Cr) in a non-breeder in a colony to 49.1 +/- 5.5 ng/mg Cr when singly housed and 21.8 +/- 2.5 ng/mg Cr when paired with a female. Plasma LH concentrations increased from 4.7 +/- 1.0 miu/ml when a non-breeder in a colony to 19.8 +/- 4.0 miu/ml when singly housed and 9.9 +/- 1.1 miu/ml when paired with a female. After pairing with a female, the pattern of urinary testosterone secretion in the male was synchronized with the ovarian cycle of the female mate, such that urinary testosterone concentrations were significantly higher during the early follicular phase of the female's cycle (P less than 0.05). These results suggest that active suppression of reproductive physiology by social cues occurs in non-breeding male naked mole-rats, and that this is readily reversible if social cues are removed and males are housed singly. When a male was subsequently paired with a female, endocrine suppression was partially reimposed on the reproductively active males, such that urinary testosterone concentrations were suppressed to values similar to those in non-breeding males, except for periods prior to mating.(ABSTRACT TRUNCATED AT 250 WORDS)     

Differences in basal energy expenditure and obesity

Objective: To assess the impact of differences in basal energy expenditure on adiposity. Research Methods and Procedures: Statistical analysis was performed on a published database giving anthropometric and energy expenditure measurements for 433 women and 335 men. Published equations derived by multiple regression analysis were used to predict basal metabolic rates in women and in men as a function of age, weight, and height. The differences between the observed and predicted rates (i.e., the residuals) were computed and expressed in terms of percentage deviation from the predicted rates of basal energy expenditure (BEE). In addition, individual body fat contents were computed using equations based on National Health and Nutrition Examination Study 3 data relating to body fat content determined by bioimpedance to BMI. Results: There is no correlation between percentage body fat content and deviations from predicted (which one would refer to as normal) BEE. Discussion: It can be concluded that relatively high or relatively low rates of BEE do not influence body weights and adiposity in a statistically identifiable manner. This contradicts and challenges the widely held view that low resting metabolic rates promote the development of obesity.