Feeding of tropical freshwater fishes: seasonality in resource availability and resource use

Summary Food resources in the environment and in the diets of small fish inhabiting two water bodies in a tropical savanna were studied during both wet and dry seasons. During the wet season (high water, abundant food) most fish species in both habitats fed predominantly on vegetation-dwelling invertebrates. Most fish species switched to alternative foods (algae and detritus) following the drastic decline in invertebrate food available towards the end of the dry season. In one habitat, this change in diet was accompanied by an increase in the volume of food intake. In the second habitat, only two larger species foraged intensively, while smaller species showed low food intake or almost ceased feeding. These differences may be explained by the high risk of predation for small fish in the second habitat. Dietary overlaps among fish species were high at the end of the dry season and moderate in the wet season. However, critical analysis of such factors as food abundance, the size and number of shared prey, and diet breadth showed that all significant overlaps were ecologically unimportant i.e. there was only weak competition for food.     

Seasonal variation in feeding habits and diet selection by wild boars in a semi-arid environment of Argentina

The wild boar, Sus scrofa, was first introduced for hunting purposes in Argentina in 1906 and presently occupies a wide range of habitats. Understanding the food habits of invasive species is important for predicting the effects of animal food consumption on the environment and on human activities, such as farming. The wild boar is an omnivorous, opportunistic species whose diet is determined by the relative abundance of different types of foods. In general, the wild boar’s diet has been widely studied in the world, both as a native and invasive species, but little is known regarding food resource selection in the Monte Desert biome. Our study assessed the seasonal variation in the diet of wild boars, as well as the nutritional quality of consumed items. Further, we determined the diet selection of this species. Diet analyses were based on faecal samples collected over two seasons (wet and dry) in 1 year. Herbs were the most frequently consumed food item, with wild boars showing a selection for them in both seasons. The wild boar uses food resources according to seasonal availability (larger trophic niche breadth under higher plant diversity, as in the wet season). In turn, within each season, it selects items of high forage quality and high carbohydrate contents. In conclusion, this foraging strategy enables wild boar to maximize energy budget through food selection in order to survive in a semi-arid environment such as the Monte Desert.     

Seasonality of reproduction by liverbearing fishes in tropical rainforest streams

Reproductive ecology, population structure, and diets of three common livebearing poeciliid fishes (Alfaro cultratus, Phallichthys amates, Poecilia gilli) from rainforest streams in Costa Rica were investigated over ten continuous months. The region experiences little annual temperature variation, and although monthly rainfall is continuous each year, two brief dry seasons typically occur. Monthly changes in indices of ovarian condition, percentages of females with developing embryos, and population size structure revealed that reproductive output by females of all three species varied seasonally. Based on testicular condition, males were reproductively active year-round, however the mean gonadal index for males of two algivorous species showed low levels of seasonal cycling that largely coincided with female variation in reproductive effort. All three species had seasonal differences in the female size-brood size relationship, whereby larger females tended to carry more embryos during the wet season. Several important adult and neonate food resources are more available in the flooded forest during the wet season, which is also the period when conspecifics and predators are at their lowest per-area densities. Three hypotheses are discussed: (1) brood size in relation to conspecific density-mating frequency, (2) reproductive allocation in response to variation in adult food resources, and (3) selection for greater reproductive effort during conditions optimal for juvenile growth and survival. Data for Alfaro were consistent with the latter two hypotheses. In Phallichthys and Poecilia, diets were poorer during wet seasons, indicating that reproductive effort does not coincide with availability of adult food resources, and that selection probably favors greater reproductive effort during periods optimal for juvenile growth and survival.     

Spatial and trophic niche differentiation in two sympatric lizards (Tropidurus torquatus and Cnemidophorus ocellifer) with different foraging tactics

Abstract In a ‘restinga’ habitat of southeastern Brazil, we studied the food habits and the microhabitat use of two lizards with distinct foraging modes: the tropidurid Tropidurus torquatus, a sit-and-wait predator, and the teiid Cnemidophorus ocellifer, a wide forager. The diet of the two species differed strongly, indicating a low level of similarity in their trophic niche. The sit-and-wait predator fed mainly on mobile prey, whereas the wide forager fed mainly on sedentary prey (larvae). The spatial niche breadth of T. torquatus was larger than that of C. ocellifer. Despite interspecific differences, the two species overlapped greatly in micro-habitat use. The data indicate that at Linhares the two lizard species differed more in food resources than in microhabitat, and that most of the food differences reflect the foraging patterns of the species.     

Effect of Seasonality on the Behavior of an Insectivorous Primate, Tarsius spectrum

The distribution and quality of food resources is generally recognized as the preeminent factor explaining much interspecific and intraspecific variation in the behavior of nonhuman primates. Primates that live in seasonal environments often show predictable responses to fluctuating resources. In order to compensate for the reduction in resource availability, primates variously switch to alternative, poorer quality food sources, increase the amount of time they spend foraging, or increase their daily path length. Some primate species reduce their group size or maximize the group dispersion. I address whether spectral tarsiers (Tarsius spectrum), which are insectivores, modify their behavior in the same ways as frugivores and folivores in response to seasonal or scarce resources. My results indicate that wild spectral tarsiers modify their activity budget in response to seasonal resources. Specifically, during periods of low resource availability, spectral tarsier males and females spent more time traveling and foraging compared to their activity budget during the wet season. Males and females not only increased the amount of time they spent foraging during times of low resource abundance but also modified their foraging behavior. During the wet season, when resource abundance was high, they consumed Orthoptera and Lepidoptera with greater frequency than during the dry season. During the dry season, when resource abundance was low, spectral tarsiers still ate numerous Orthoptera and Lepidoptera, but they also increased consumption of Coleoptera and Hymenoptera. Spectral tarsiers were also more likely to be involved in territorial disputes during the dry season than during the wet season. Intragroup encounters decreased in frequency in the dry season versus the frequency of encounters during the wet season.     

Bats, flowers and fruit: evolutionary relationships in the Old World

Old World phytophagous bats (Megachiroptera: Pteropodidae) number 173 species of which 79% are Asian and 21% African. Bats arose, presumably monophyletically, in the early Tertiary, the Megachiroptera soon diverging from the Microchiroptera. By the Cretaceous-Tertiary boundary the major groups of modern angiosperms were present, some of these probably being pollinated nocturnally by large insects and non-flying mammals and others with seeds dispersed by terrestrial vertebrates. Early bats were perhaps initially attracted to such flowers and fruit by the insects found around them, later finding the plants themselves nutritious. Megabats today feed upon floral resources, fruit and leaves from a total of at least 188 plant genera in 64 families. They may effect both pollination and seed-dispersal, and both bat-flower and bat-fruit syndromes are commonly recognized. Individual species are generally catholic in their feeding, favoured food varying with locality and season. Depending upon roosting habits and season, megabats may travel considerable distances each night to feed and may undertake seasonal migrations. Their feeding in orchards may sometimes require their control, but the future of certain species is more seriously threatened by slaughter for food and particularly by habitat destruction.     

Comparative ecology of three pteropid bats in Rio Muni, West Africa

Data on morphologic features, habits, stratification, flight, and thermal response are presented for Eidolon helvum, Epomops franqueti , and Micropteropus pusillus from Rio Muni, West Africa. Relative abundance and reproduction activities of these pteropids are concordant with seasonal rainfall and availability of foods in Rio Muni. There are major differences between these sympatric species with regard to roosting sites, foods and feeding behaviour, levels of flight, and aerodynamic properties. Thermal responses vary between the species and are correlated with differences in behavioural thermoregulation.     

Diet and habitat overlap in two sympatric primate species, the Tana crested mangabey Cercocebus galeritus and yellow baboon Papio cynocephalus

Diet and habitat overlap was studied in two sympatric primate species sharing two neighbouring patches of fragmented gallery forest in Tana River, Kenya. Systematic data on feeding and ranging behaviour was collected on one group each of the Tana crested mangabey Cercocebus galeritus and yellow baboon Papio cynocephalus between August 1992 and February 1993. When rainfall was low and fruit resources scarce, yellow baboons spent most of their foraging time in the open woodlands while mangabeys foraged within the forest. At this period, diet and habitat overlaps between the two species were low. As rainfall increased, followed by a gradual rise in fruit availability, yellow baboons shifted their foraging range and both species became confined to the forest habitat. Consequently, both diet and habitat overlaps increased, peaking at the end of the rainy season. Mangabeys showed a reduced within-group dispersal and also spent significantly less time foraging in a given forest patch when yellow baboons were also present within the same forest patch.
Increased habitat and diet overlaps during the wet season need not have resulted in increased interspecific competition for food because at this period, fruits were readily available in the forest.     

Reproduction and growth in three species of West African fruit bats

This study outlines the reproduction periods of the African fruit bats Epomops buettikoferi, Epomophorus garnbianus , and Micropteropus pusillus at two West African savanna sites: a wet southern Guinea savanna and a drier southern Sudanese savanna. At both sites the two annual birth periods were timed such that both lactation by only one weaning period coincided with the rainy season peaks in fruit availability. On this basis we propose that lactation rather than weaning was the important determinant of the timing of reproduction. There was no evidence of cycling of testes size in males corresponding with the seasonal mating periods. E. buettikoferi and M. pusillus females mated at six months and gave birth at the age of 12 months. Males of the two species reached puberty by 11 months and seven months, respectively. Growth rates did not differ between cohorts growing through the wet and dry season and E. buettikoferi and M. pusillus grew at rates of 276 mg/day and 116 mg/day, respectively.     

Resource partitioning among three woodpecker species Dendrocopos spp. during the breeding season

Food composition, prey size utilization and foraging behaviour of three sympatric woodpecker species ( Dendrocopos major, D. medius, D. minor ) were studied in an oak forest near Budapest during the breeding season in 1983 and 1984. Considering these three aspects of feeding, the great spotted woodpecker is a generalist species. Food composition of this species resembled the arthropod supply on the bark of trees more than those of the other two species. The bark of the trees seems to be a relatively unproductive microhabitat in the breeding season, so woodpecker species use, to different degrees, the food supply of the foliage as well. The food and the foraging behaviour of the middle spotted woodpecker show that this species feeds on prey living both on barks and in the foliage; it occupies-an intermediate position between the great and the lesser spotted woodpeckers. Prey size did not correlate with predator size suggesting that woodpeckers adapted not to the summer resources but rather the winter ones.     

How equally sized piscivorous birds and fish sharing common food resources may reduce possible feeding interactions between them?

The diets of sympatric predators may overlap, especially when their body sizes are similar and foraging area is relatively small. It may be also supposed that some differences in their foraging strategies may counteract competitive interactions among them, and therefore be of advantage to these species. To reveal such phenomena the composition of food of cormorant and adult pikeperch was studied in the Dobczyce Reservoir (S Poland) from June to November 2002. The main prey species were the same and the range of prey size was similar for both piscivores. Despite these similarities, the potential for dietary overlap was strongly reduced due to two differences in their foraging patterns: (1) different preferred prey species (cormorants foraged mainly roach, whereas pikeperch selected juvenile percids); (2) different size of simultaneously selected prey (in summer, cormorants selected larger prey, while in autumn larger prey was selected by pikeperch). These differences may be explained by some general features of birds and fishes, which determine the costs to the individual of capturing prey. The observed selection of different prey species and sizes may be also important for the co-occurrence of other piscivorous birds and fishes sharing common food resources.     

Ranging behaviour and activity of red foxes (Vulpes vulpes: Mammalia) in relation to environmental variables, in a Mediterranean mixed pinewood

The relationships between the main abiotic environmental variables, food availability, dict, activity and home range size of the fox in a Mediterranean mixed pinewood were assessed. Food habits were determined by faecal analysis on a weekly basis. Food availability was measured by weekly censuses. Radio-tracking allowed the detection of fox movements and activity. Individual differences shown by a sample of four radio-tagged foxes were analysed. Juniper berries (Juniperus oxycedrus) were the staple of the diet during the year, except in summer. Insects, mainly nocturnal Orthoptera and cicada larvae (Cicada orni), were the main prey in the warm season. Large vertebrates, mostly wild boars (Sus scrofa), were scavenged in winter. In the warm season foxes' activity depended largely on the pattern of activity of prey. Food habits tended to depend on food availability, which, in turn was mainly influenced by temperature. In spite of habitat similarity, individual differences in food habits were detected within the same season. Home ranges were larger in pinewood than in neighbouring heterogeneous habitats. Individual home range sizes were: (i) directly correlated to Orthoptera abundance and distribution; (ii) inversely correlated to the abundance of cicada larvae in diet. Resident males may exist in fox society, exploiting other males' territories when good clumped food resources are available.     

Patterns of roost site selection and use by Southern Ground-Hornbills in north-eastern South Africa

Different habitats may be used for the needs of various aspects of an animal’s life. Southern Ground-Hornbill Bucorvus leadbeateri groups announce their presence within year-round territories by calling at dawn from their overnight roost sites. Knowledge on ground-hornbill roosting habits is limited. Groups roost in large trees, apparently close to where they end up after daily foraging. We investigated patterns of roost site selection and use for four Southern Ground-Hornbill groups in the Associated Private Nature Reserves, north-eastern South Africa, based on data from GPS-satellite transmitters. The number of roost sites used per month averaged 15.4 ± 4.7 across all groups, indicating little evidence of strong preferences for specific sites. This number was least when groups were breeding, decreasing throughout the early wet season (October–December) and was lowest during the late wet season (January–March) when actively breeding groups frequently roosted close to the nest (54–83% of roosts <1 000 m of the nest). As might be expected, the mean monthly number of nights per roost peaked during the breeding season (December–January). Riparian habitats were preferred for roosting during the breeding season, whereas disturbed areas, as well as Combretumand mopane-dominated habitats were preferred during the dry non-breeding season. Adequate large trees not only for nesting, but also for roosting, particularly in riparian habitats, may therefore be an important and potentially limiting factor for the successful reproduction of Southern Ground-Hornbills.     

Environmental constraints on the foraging behaviour of a dwarf antelope (Madoqua kirkii)

Dik-diks (Madoqua sp.) inhabit semi-arid regions and experience very different conditions of food availability and quality between wet and dry seasons. By comparing the behaviour of dik-diks between these two seasons, we identified environmental constraints affecting their feeding strategies. In both seasons foraging time was limited by high mid day temperatures. In the wet season a high intake rate compensates for the loss in foraging time, but in the dry season water and protein become limiting. To meet minimum daily water requirements in the dry season dik-diks fed on plant species that they avoided during the wet season. Analysis at the plant species level showed higher species selectivity in the wet season than in the dry season. In a multiple regression analysis food species preferences were best explained by relative abundance and water content in the dry season, and by dry matter content in the wet season. In the wet season the daily dry-matter intake of dik-diks in the field was only about 10% higher than the theoretically predicted minimum for a ruminant of this body weight, while protein and water intake were about 3 times as high. This suggests that the most limiting dietary component in the wet season is energy. In the dry season the daily intake of all dietary components is lower than the theoretical minimum required, and also lower than the values suggested by laboratory studies of dik-diks. This dry season deficit is presumably met from body reserves. Dry season water intake was approximately 30% of the intake observed in laboratory studies indicating that dikdiks are even better adapted to arid conditions than suggested by physiological experiments.     

Ecological determinants of herd size in the Thornicroft’s giraffe of Zambia

Ecological factors have a pervasive impact on animal population sizes and the structure of their social systems. In a number of ungulate species, predator pressure exerts a major influence on group size. Given that giraffe (Giraffa camelopardalis) live in an extremely flexible social system, and that breeding is nonseasonal, they are an ideal species for examining how ecological variables contribute to fluctuations in herd size. We present an analysis of 34 years of data on a population of Thornicroft’s giraffe (G. c. thornicrofti Lydekker 1911) that reveal how herd size changes with season and habitat. Sex differences in herd size were apparent, with bulls often travelling as singletons, whereas cows were generally observed with conspecifics. Herds were larger during the wet than dry season, but herd size changed in a parallel fashion across habitats. Giraffe herds were smaller in woodland and thicket areas than in open habitats, regardless of season. We suggest that the regular fluctuations in herd size among giraffe indicate a fission/fusion social system embedded within a larger social community. We conclude that changes in herd size among giraffe reflect a dynamic process regulated by individuals adjusting the number of associates based upon an interaction of foraging, reproductive, social and antipredator strategies.     

OBSERVATIONS ON THE FOOD HABITS OF TAME WARTHOG AND ANTELOPE IN UGANDA

Monthly observations were made of the food habits of tame warthog, reedbuck and kob over a period of a year. Measurements were taken of the shoulder height and size of mouth of the animals at different stages of their growth. Whilst a few species of grass were regularly preferred, the majority showed fluctuations in their preference. Some consistent seasonal differences were apparent. Most herbs were avoided. Large differences in food habits occurred between warthog and antelope; however, there was some overlap during the rains when food was abundant and at the height of the dry season when it was scarce. Some differences were apparent between kob and reedbuck and between individual warthog and reedbuck. Intraspecific differences, however, were less noticeable than interspecific differences. The height of the animal and the size of its mouth were important in determining the portion of the sward available for food and the degree of selection possible within the available food supply.     

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.     

Water depth selection, daily feeding routines and diets of waterbirds in coastal lagoons in Ghana

Water depth requirements, diet, feeding styles and diurnal activity patterns are described for waterbirds using two brackish water lagoon systems in coastal Ghana, the Songor and Keta Lagoons. We project the habitat and activity data on a guild structure defined on the basis of individual feeding style and the sensory mechanism used to detect food. A total of 3199 flocks containing 118,648 individuals of 36 different waterbird species were examined during October-November 1994. Feeding habitats varied from dry mudflats to wet mud and shallow water of not more than 20 cm. The depth of water selected by waterbirds for foraging (but not for roosting) was correlated with tarsus length. Foraging birds exhibited a wide range of feeding styles using visual and/or tactile means for detecting prey: pecking, probing, stabbing, sweeping and ploughing, sometimes feeding singly, communally or socially in loose or dense flocks. Prey items taken ranged from seeds of Widgeongrass Ruppia maritima to invertebrates (mainly polychaetes, molluscs and crabs) and fish, mainly juvenile Tilapia. The daytime was spent on two main activities, feeding and roosting, with a small fraction of the time (average of 10% for 25 species) spent on comfort activities. The waterbirds exhibited either a circadian (most waders, except Common Sandpipers Actitis hypoleucos and Turnstones Arenaria interpres) or a diurnal foraging activity pattern (herons and terns), with no purely nocturnal species. Some species fed throughout the day, others showed peak foraging at various times of the day. The proportion of time spent foraging was related to guild (highest in visual and tactile surfaceforaging waders) and was negatively correlated with the size of the species. We conclude that the observed patterns in the use of the 24-h day by waterbirds for foraging are not species specific but vary depending on conditions on the feeding grounds. Nocturnal foraging is a normal and a regular strategy used by waterbirds to obtain enough food to fulfill their energetic requirements, so that irrespective of the sensory mechanism used to detect prey and the conditions prevailing on the feeding grounds, waterbirds forage day and night as dictated by their energetic needs. Water depth appears to be the key environmental factor controlling the availability of food for the waterbirds in the Ghanaian lagoons.     

Plant food resources and the diet of a parrot community in a gallery forest of the southern Pantanal (Brazil).

Neotropical parrots usually forage in forest canopies for nectar, flowers, leaves, fruit pulp, and seeds. As they have no all-purpose territories, these birds usually exploit vegetation mosaics in order to use plentiful resources as they become available. In this study we examine the use of a gallery forest in the southern Pantanal (Brazil) by a diverse parrot community that ranged from Brotogeris chiriri (a small species) to Ara chloroptera (a large one). Plant food resources principally used by parrots were abundantly available during the rainy season (fleshy fruits), the annual floods (fleshy fruits), and the dry season (flowers). While both smaller and larger species foraged on fruits, parakeets largely consumed the pulp, while larger parrot species used pulp and seeds. In the dry season parakeets foraged extensively on nectar, especially Inga vera nectar that was abundantly available during the last two months of the dry season, the harshest period of the year. Among larger parrots, only Propyrrhura auricollis frequently harvested nectar. Fruits maturing during floods, despite being fish- or water- dispersed were extensively used by the parrots. Hence, unlike what happens in most other Neotropical dry forests, occurrence of a fruiting peak during the annual flooding, which occurs in the transition from the wet to the dry season, constitutes an extra and significant episode of food availability, since in this period, fruit production normally declines. Therefore, the unique and abundant availability of flowers and fruits in this gallery forest may account for the presence of large parrot populations in the southern Pantanal.     

Emergence of complexity in evolving niche-model food webs

We have analysed mechanisms that promote the emergence of complex structures in evolving model food webs. The niche model is used to determine predator-prey relationships. Complexity is measured by species richness as well as trophic level structure and link density. Adaptive dynamics that allow predators to concentrate on the prey species they are best adapted to lead to a strong increase in species number but have only a small effect on the number and relative occupancy of trophic levels. The density of active links also remains small but a high number of potential links allows the network to adjust to changes in the species composition (emergence and extinction of species). Incorporating effects of body size on individual metabolism leads to a more complex trophic level structure: both the maximum and the average trophic level increase. So does the density of active links. Taking body size effects into consideration does not have a measurable influence on species richness. If species are allowed to adjust their foraging behaviour, the complexity of the evolving networks can also be influenced by the size of the external resources. The larger the resources, the larger and more complex is the food web it can sustain. Body size effects and increasing resources do not change size and the simple structure of the evolving networks if adaptive foraging is prohibited. This leads to the conclusion that in the framework of the niche model adaptive foraging is a necessary but not sufficient condition for the emergence of complex networks. It is found that despite the stabilising effect of foraging adaptation the system displays elements of self-organised critical behaviour.