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.     

Temporal patterning of feeding by hummingbirds

For five species of hummingbirds in the laboratory, time between meals was related to energy intake on the first meal and rate of energy expenditure between meals. Field observations gave similar results. Average meal sizes were similar at one intake rate independent of food caloric density; females averaged longer bouts than males. When rate of intake was approximately halved, meal duration approximately doubled and volume intake remained similar. We postulate that feeding is initiated when crop contents reach a lower threshold and that feeding is terminated after ingestion of an optimal volume determined by the added weight of the meal.     

Energy,predators and the behavior of feeding hummingbirds

Summary Current evidence suggests that many animals trade off energy gain against the risk of predation while feeding. In contrast, energetic considerations alone have proven successful in explaining and predicting the behaviour of feeding hummingbirds. This success may reflect the relative lack of natural predation on hummingbirds, but this study suggests that it may additionally reflect the lack of studies taking a predation perspective. In particular, when Anna's hummingbirds (Calypte anna) are faced with an obstructed view of their surroundings, they engage in behaviour suggestive of anti-predatory vigilance. In doing so, they voluntarily reduce their rate of energy intake. These birds also forgo better feeding opportunities that occur close to the ground, where observations suggest they are wary of opportunistic predators such as roadrunners (Geococcys californianus). While energy-based concepts will remain useful in the study of hummingbird feeding behaviour, the lack of predation on these birds should not be equated with an insensitivity to the risk of predation. This realization may lead to further insights into hummingbird-plant interactions, and hummingbird biology in general.     

Epigenetic regulation of trophic morphology through feeding behaviour in Arctic charr, Salvelinus alpinus

Several models of speciation suggest that in species that are phenotypically plastic, selection can act on phenotypic variation that is environmentally induced in the earliest stages of divergence. One trait that could be subject to this process is foraging behaviour, where discrete foraging strategies are common. One species which is highly plastic in the expression of phenotype, the Arctic charr, Salvelinus alpinus (L.), is characterized by discrete variation in the anatomy of the head and mouthparts. These traits have been shown to have a functional significance, but the expression of which is thought to be at least partly phenotypically plastic. Here we test the hypothesis that foraging behaviour may regulate the anatomy of the head and mouthparts in Arctic charr. In a dyad experiment, size‐matched pairs of fish from a mixed family group were fed a diet of either Mysis (a hard‐bodied shrimp) or Chironomid larvae. Nine morphometric measures of head dimensions that describe wild trophic morphs were measured at the start of the experiment and 24 weeks later. Principal component scores of size‐corrected morphometric measures showed highly significant differences between fish exposed to the two diets. Univariate ANOVA analysis of the head morphometric variables showed that fish fed on Chironomids developed longer, wider jaws, longer heads and a larger eye for a given body length than did those fish fed upon Mysis. We conclude that foraging anatomy in Arctic charr is phenotypically plastic and that variation in foraging behaviour that results in feeding specialization in the wild could induce variation in head anatomy. This in turn could reinforce foraging specialization. Very rapid epigenetic divergence into distinct feeding morphs (as demonstrated here) would allow selection to act at more than one mode and thus could promote rapid evolutionary divergence, initially prior to genetic segregation, in species which are highly plastic. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78 , 43–49.     

Effect of sexual dimorphism in bill length on foraging behavior: an experimental analysis of hummingbirds

We examined whether sexual differences in trophic morphology are associated with sexual differences in foraging behavior through two laboratory experiments on rufous hummingbirds (Selasphorus rufus) designed to compare probing abilities (maximum extraction depths) and handling times of sexes at flowers. Bills of female S. rufus are about 10.5% longer than bills of males, and this difference was associated with sexual differences in foraging abilities. Maximum extraction depths of female S. rufus were significantly greater than those of males, and no overlap between the sexes was observed. Moreover, handling times of females were shorter than handling times of males at flowers having longer corollas (15 mm). Thus, because of their longer bills, female S. rufus have the potential to feed from longer flowers than males, and can do so more quickly. We suggest that no single mechanism is responsible for the evolution of sexual dimorphism in bill lengths of hummingbirds, but rather that the dimorphism probably reflects the combined effects of reproductive role division and intersexual food competition, and possibly, sexual selection.     

Estimation of social genetic effects on feeding behaviour and production traits in pigs

Pigs are housed in groups during the test period. Social effects between pen mates may affect average daily gain (ADG), backfat thickness (BF), feed conversion rate (FCR), and the feeding behaviour traits of pigs sharing the same pen. The aim of our study was to estimate the genetic parameters of feeding behaviour and production traits with statistical models that include social genetic effects (SGEs). The data contained 3075 Finnish Yorkshire, 3351 Finnish Landrace, and 968 F1-crossbred pigs. Feeding behaviour traits were measured as the number of visits per day (NVD), time spent in feeding per day (TPD), daily feed intake (DFI), time spent in feeding per visit (TPV), feed intake per visit (FPV), and feed intake rate (FR). The test period was divided into five periods of 20 days. The number of pigs per pen varied from 8 to 12. Two model approaches were tested, i.e. a fixed group size model and a variable group size model. For the fixed group size model, eight random pigs per pen were included in the analysis, while all pigs in a pen were included for the variable group size model. The linear mixed-effects model included sex, breed, and herd*year*season as fixed effects and group (batch*pen), litter, the animal itself (direct genetic effect (DGE)), and pen mates (SGEs) as random effects. For feeding behaviour traits, estimates of the total heritable variation (T² ± SE) and classical heritability (h² ± SE, values given in brackets) from the variable group size model (e.g. period 1) were 0.34 ± 0.13 (0.30 ± 0.04) for NVD, 0.41 ± 0.10 (0.37 ± 0.04) for TPD, 0.40 ± 0.15 (0.14 ± 0.03) for DFI, 0.53 ± 0.15 (0.28 ± 0.04) for TPV, 0.66 ± 0.17 (0.28 ± 0.04) for FPV, and 0.29 ± 0.13 (0.22 ± 0.03) for FR. The effect of social interaction was minimal for ADG (T² = 0.29 ± 0.11 and h² = 0.29 ± 0.04), BF (T² = 0.48 ± 0.12 and h² = 0.38 ± 0.07), and FCR (T² = 0.37 ± 0.12 and h² = 0.29 ± 0.04) using the variable group size model. In conclusion, the results indicate that social interactions have a considerable indirect genetic effect on the feeding behaviour and FCR of pigs but not on ADG and BF.     

Host-associated variation in sexual size dimorphism and fitness effects of adult feeding in a bruchid beetle

We studied the effect of larval host (two Convolvulus L. species, Convolvulaceae) on sexual size dimorphism and on the fitness consequences of adult feeding in the bruchid beetle Megacerus eulophus (Erichson) (Coleoptera: Bruchidae). Whereas Convolvulus chilensis Pers. occurs in low density in semiarid habitats, Convolvulus bonariensis Cav. occurs in less stressful environments and exhibits higher population density. Host plants neither differ in seed mass nor in seed nitrogen content, and there were no consistent host‐associated differences in female fitness. Consequently, host quality was considered to be similar. Sexual size dimorphism, expressed as the female:male ratio of body size, was significantly greater in C. chilensis than in C. bonariensis. A greater female size in the C. chilensis population could be selectively advantageous due to higher survival during host‐plant search in stressful environments. Female longevity was affected by food availability (starvation vs. honey‐pollen solution) and by the interaction between food availability and larval host. Fed females lived longer than starved females. This trend was found in both hosts, but it was of greater magnitude in C. chilensis than in C. bonariensis. Fecundity was significantly affected by adult feeding and larval host, and marginally affected by their interaction. Fed females laid more eggs than starved females. However, the increase in fitness of fed females was greater in C. chilensis. Results might be explained by differential selection on plasticity in life‐history traits in contrasting environments. Females in stressful environments should take a greater advantage of the transient availability of resources.     

Seed dimorphism and germination behaviour in Salicornia patula

One of the annual species of the genus Salicornia from the Mediterranean coast, Salicornia patula presents a seed dimorphism. This morphological seed dimorphism is linked with a physiological ‘dimorphism’ with regard to some requirements for germination. Central seeds, in the inflorescence, are larger, are dispersed attached to the perianth, have no light requirement for germination and seem to have a low germination response to salinity. The lateral seeds, which are free from perianth envelopes, require light and cold pretreatment and seem to have a high sensitivity to salinity. Salicornia patula colonizes unstable transition zones between permanently flooded muds and perennial vegetation. These zones have a winter flooding period. The beginning of the flooding period and of the dry period may vary from year to year, depending on the erratic distribution of autumn and winter rains on the Mediterranean coast. So, in some years, a seedling population may be destroyed before reaching the stage of seed production. Seed dimorphism might cause germination to the more distributed in time, thus reducing the chances of extinction of a complete generation. In the present paper salinity and temperature response are described more precisely, are investigated, some aspects of the longevity of the two types of seed, and the adaptative significance of the corresponding dispersal and germination syndromes are discussed in more detail.     

The effects of feeding and housing on the behaviour of the laboratory rabbit

The effects of housing, feeding time and diet composition on the behaviour of the laboratory rabbit were examined. The animals were caged individually in single or double metal cages with perforated metal floors, metal walls, and bars in the front, or kept as a group in floor pens. The light/dark cycle was 12/12 h with light from 04:00 to 16:00 h and 30 min twilight. One experiment compared feeding equal energy levels of a high energy diet (10.1 MJ/kg) and with a low energy diet (7.0 MJ/kg) at 08:00 h. The second experiment compared feeding the high energy diet at 08:00 h and at 14:00 h. In both studies the behaviour of the rabbits was recorded between 08:00 and 14:00 h and between 16:00 and 22:00 h. Feeding the animals at 14:00 h reduced abnormal behaviour during the dark period compared to feeding at 08:00 h, whereas no difference in behaviour could be detected between feeding a high-energy and a low-energy diet at 08:00 h. Animals in floor pens generally showed less abnormal behaviour than caged animals. The results indicate that the welfare for caged rabbits can be improved by feeding the animals in the afternoon rather than in the morning.     

Dairy intensification in developing countries: effects of market quality on farm-level feeding and breeding practices

Smallholder dairy production represents a promising income generating activity for poor farmers in the developing world. Because of the perishable nature of milk, marketing arrangements for collection, distribution and sale are important for enhanced livelihoods in the smallholder dairy sector. In this study we examined the relationship between market quality and basic feeding and breeding practices at farm level. We define market quality as the attractiveness and reliability of procurement channels and associated input supply arrangements. We took as our study countries, India with its well-developed smallholder dairy sector, and Ethiopia where the smallholder dairy industry has remained relatively undeveloped despite decades of development effort. We conducted village surveys among producer groups in 90 villages across three States in India and two Regions in Ethiopia. Producer groups were stratified according to three levels of market quality – high, medium and low. Data showed that diet composition was relatively similar in India and Ethiopia with crop residues forming the major share of the diet. Concentrate feeding tended to be more prominent in high market quality sites. Herd composition changed with market quality with more dairy (exotic) cross-bred animals in high market quality sites in both India and Ethiopia. Cross-bred animals were generally more prominent in India than Ethiopia. Herd performance within breed did not change a great deal along the market quality gradient. Parameters such as calving interval and milk yield were relatively insensitive to market quality. Insemination of cross-bred cows was predominantly by artificial insemination (AI) in India and accounted for around half of cross-bred cow inseminations in Ethiopia. Data on perceptions of change over the last decade indicated that per herd and per cow productivity are both increasing in high market quality sites with a more mixed picture in medium and low-quality sites. Similarly dairy-derived income is on the increase in high market quality sites. This is accompanied by a strong increase in stall feeding at the expense of grazing. The study indicates that the first constraint to intensification of dairy production in Ethiopia is the genetic quality of the herd. There is less scope for improved AI provision in India since the cross-bred herd is mainly serviced by AI already. However, as for Ethiopia, there is considerable scope for closing yield gaps in India through improved feed use and supply. Results strongly show that well-developed markets with good procurement arrangements are key for sustainable dairy intensification.     

Permethrin-impregnated bednet effects on resting and feeding behaviour of lymphatic filariasis vector mosquitoes in Kenya

The impact of permethrin-impregnated bednets on resting and feeding behaviour of mosquito vectors of Wuchereria bancrofti, causing human lymphatic filariasis was studied in six pairs of villages (treated and untreated) before and after intervention. The study villages were in Kwale District, near the coast of Kenya, where Bancroftian filariasis is highly endemic, transmitted by a combination of both anopheline and culicine mosquito vectors. Mosquitoes were collected weekly in each village, indoors (using pyrethrum spray catches) and outdoors (using pit traps) during 3–4 months following the long rainy season. Of the filariasis vector species of mosquitoes collected in 1994 before intervention, 33.6% were members of the Anopheles gambiae complex, 30% were An. funestus and 36.4% were Culex quinquefasciatus. PCR analysis of the An. gambiae complex species collected in 1995 demonstrated that 98.5% were An. gambiae sensu stricto, 1% An. arabiensis and 0.5% An. merus. Introduction of impregnated bednets in 1995 significantly reduced the number of indoor-resting An. gambiae s.l. by 94.6% and An. funestus by 96.7%, but there was no change in the number of Cx quinquefasciatus collected indoors. The number of outdoor-resting An. gambiae s.l. was significantly reduced, whereas densities of An. funestus and Cx quinquefasciatus remained unaffected outdoors. ELISA analysis of mosquito bloodmeals showed a shift from human to animal feeding after the introduction of treated nets. The human blood index (HBI) for indoor resting Cx quinquefasciatus was reduced from 93.1% to 14.4%. Vector potential based on the HBI and mosquito density was estimated to be reduced by 99% for An. gambiae s.l., 98% for An. funestus and 97% for Cx quinquefasciatus and vectorial capacity would be suppressed even more by the impact on the vector survival rates (not measured). These results suggest that permethrin-impregnated bednets give effective personal protection against transmission of W. bancrofti by An. gambiae, An. funestus and Cx quinquefasciatus in East Africa.     

Foraging behaviour of planktivorous fish in artificial vegetation: the effects on swimming and feeding

In the littoral zones of lakes, aquatic macrophytes produce considerable structural variation that can provide protection to prey communities by hindering predator foraging activity. The swimming and feeding behaviour of a planktivore, Pseudorasbora parva(Cyprinidae) on its prey (Daphnia pulex) was studied in a series of laboratory experiments with varying densities (0, 350, 700, 1400, 2100 and 2800 stems m^–2) of simulated submerged vegetation. Prey availability was varied from 0.5, 1.0, 2.0, 5.0, 10.0 and 25.0 prey l^–1. As the stem density increased, the predator's swimming speed and the number of prey captured decreased relative to feeding in open water. A good relation existed between the number of successful prey captures and swimming speed with the average stem distance to fish body length ratio (D). An abrupt reduction in feeding and swimming was recorded when D was reduced to values less than one.     

Variations in feeding and social behaviour in a captive breeding group of harbour seals (Phoca vitulina)

Most Phocids are thought to reduce their food intake during the breeding and moulting seasons, even though fish are not in short supply. The present study of a captive breeding colony of harbour seals explored the relationship between changes in amount eaten, social behaviour and general activity levels. Even though the animals were given free access to food, over the year the daily amount they ate varied in three cycles. Food eaten fell below average between pupping and mating in June‐July, and fell again in November until early December, for a third time in mid February, and again in early May. Levels surpassed average during and shortly after moulting, and again in late December. Observations made during the summer and early fall revealed that the adult males became more active, and engaged in more social interaction when they were eating less. The female showed no such activity change as a function of appetite. Play occurred primarily when food levels were high. These results are discussed in the context of systems interaction theories of motivation.     

尖吻蝮人工养殖灌喂技术研究

目的探讨采用灌喂器人工灌喂尖吻蝮技术的可行性。方法采用湖南京湘源蛇类养殖有限公司自主研制的灌喂器人工灌喂尖吻蝮,观察记录随机抽取的30条尖吻蝮幼蛇及30条尖吻蝮成蛇在机器灌喂后每条幼蛇的体重数据,每3个月测量1次,观察12个月统计尖吻蝮的体重增长情况,并与常规灌喂方法饲养的尖吻蝮进行比较。结果灌喂器饲养的尖吻蝮幼蛇成活率为86.67%,平均体重达（482.39±40.19）g;灌喂器饲养的尖吻蝮成蛇成活率为96.67%,平均体重达（1346.13±117.51）g。而同期常规灌喂技术饲养的尖吻蝮幼蛇全部死亡,尖吻蝮成蛇成活率为76.67%,平均体重（878.56±82.39）g。结论人工灌喂器饲养的尖吻蝮幼蛇及成蛇的体重增长快速,成活率高,值得推广。     

Intraspecific differences in behaviour and physiology: effects of captive breeding on patterns of torpor in feathertail gliders

Studies on the physiology of mammals and birds are often conducted using captive-bred individuals and it is commonly assumed that the resulting data are representative of individuals living in the field. To investigate whether these assumptions are justified, we quantified morphological, behavioural, and physiological variables of the small marsupial feathertail glider (Acrobates pygmaeus). We compared three populations: (i) individuals from a cool-temperate, montane area, (ii) individuals form a subtropical, coastal area, and (iii) captive-bred individuals. Captive-bred gliders differed from the montane field gliders in morphology (longer tails and snouts), behaviour (longer activity periods) and physiology (less frequent torpor, shorter torpor, shallower torpor, higher metabolic rates during rest and torpor, and slower rates of rewarming). Most of these differences were also apparent between the captive-bred and the coastal field gliders. Unlike both field populations, captive-bred gliders often became hypothermic and were unable to rewarm. In contrast to the other physiological variables, the minimum body temperatures defended during torpor and the corresponding air temperatures differed between the montane and coastal field gliders, but were similar in coastal field and captive-bred gliders. Our study shows that morphology, behaviour and physiology can be strongly affected by breeding in or acclimation to captivity. The poor expression of torpor and thermal performance of the captive-bred gliders raises the question of whether they possess the physiological capability for survival in the wild. Even though captive breeding appears to have only minor effects on some physiological variables, data from captive-bred individuals should only be extrapolated to the field with caution.     

The effects of light intensity and algae-induced turbidity on feeding behaviour of larval striped trumpeter

Striped trumpeter larvae reared in algal cell‐induced turbid water (greenwater) fed equally well in clearwater in a light intensity range of 1–10 μmol s^‐1 m^‐2, when evaluated in terms of both the proportion of larvae feeding and larval feeding intensity. An ontogenetic improvement in photopic visual sensitivity of larvae was indicated by improved feeding at 0·1 μmol s^‐1 m^‐2, from 26±5% of larvae feeding and 0·027±0·005 rotifers consumed per feeding larva min^‐1 on day 8, to 96±2% and 0·221±0·007 rotifers consumed larva^‐1 min^‐1 on day 23 post‐hatching. Algal cell‐induced turbidity was shown to reduce incident irradiance with depth, indicated by increasing coefficients of attenuation (1·4–33·1) with increasing cell densities (0–2×10⁶ cells ml^‐1), though light intensities in the feeding experiment test chambers, at the algal cell densities tested, were within the optimal range for feeding (1–10 μmol s^‐1 m^‐2). Algae‐induced turbidity had different effects on larval feeding response dependent upon the previous visual environment of the larvae. Young larvae (day 9 post‐hatching) reared in clearwater showed decreased feeding capabilities with increasing turbidity, from 98±1% feeding and 0·153±0·022 rotifers consumed larva^‐1 min^‐1 in clearwater to 61±10% feeding and 0·042±0·004 rotifers consumed larva^‐1 min^‐1 at 56 NTU, while older clearwater reared larvae fed well at all turbidities tested. Likewise, greenwater reared larvae had increased feeding capabilities in the highest algal cell densities tested (32 and 66 NTU) compared with those in low algal cell density (6 NTU), and clearwater (0·7 NTU) to which they were naïve.