Influence of the abundance of food resources on the feeding habits of the red fox, Vulpes vulpes, in western Switzerland

J.-M. WEBER
The influence of food abundance on red fox feeding habits was studied in the Swiss Jura Mountains between November 1990 and February 1992, during a low density period in the water vole population cycle. Six food categories–water voles, small mammals, domestic stock, earthworms, wild fruits and exploitable scraps–were censused during the present research and their seasonal abundances were estimated in fixed plots. Fox scats were collected and analysed at regular monthly intervals.
The diet of foxes was found to be correlated with seasonal food supply. A positive significant correlation between seasonal abundances and relative proportions in the diet was found for food categories showing a clear seasonality (earthworms, wild fruits and exploitable scraps), whereas no correlation was recorded for the other food resources. Diet composition observed in the present study was compared with that recorded in the same study area between 1988 and 1990 by Weber & Aubry (1993), when water voles were very common and accounted for more than 50% of the diet. A significant change was recorded in red fox feeding habits: the most important aspects of this modification of the diet was the decrease in water vole importance accompanied by the increase in the frequency of occurrence of other food categories, such as wild fruits and scavengeable items.     

Influence of fruit availability on macronutrient and energy intake by female chimpanzees

Daily energy intake of adult female mammals is influenced by environmental conditions and physiological requirements, including reproduction. We examined the effects of fruit availability on macronutrient and metabolisable energy (ME) intake by adult female chimpanzees (Pan troglodytes schweinfurthii) of the Kanyawara community in Kibale National Park, Uganda, from January 2014 through June 2015. Drupe fruits were abundant for 4 months, whereas the other 14 months were dominated by fig fruits. The mean daily intake of food (dry matter) and ME did not differ between drupe‐months and fig‐months. However, foraging costs were higher during fig‐months, as indicated by a 20% increase in feeding time. Furthermore, during drupe‐months female chimpanzees ingested more water‐soluble carbohydrates and lipids, and less available protein and neutral detergent fibre. Although ME intake did not differ consistently between drupe‐months and fig‐months, they consumed more on days when ripe fruit dominated the diet than when leaves and pithy stems dominated the diet. Our data suggest that differences in diet quality between drupes and figs can have important effects on frugivore foraging and that they influence net energy gain more by their effects on macronutrient composition or foraging cost than by their direct impact on energy intake.     

Diet of the gracile mouse opossum (Gracilinanus microtarsus) (Didelphimorphia: Didelphidae) in a Brazilian cerrado: patterns of food consumption and intrapopulation variation

The diet of the gracile mouse opossum Gracilinanus microtarsus was studied in a cerrado remnant in south-eastern Brazil through the analysis of faeces sampled from adult individuals. Patterns of food resource consumption were assessed using the statistics of per cent occurrence. Intrapopulation variation in the number of food items detected in faeces as a function of relevant factors was inferred using generalized linear models. The latter statistical formalism also allowed variation in the number of food items detected in faeces to be interpreted in terms of rate ratios among levels of significant factors. Insects, spiders, snails and fruits were detected in the faeces of G. microtarsus , with insects, particularly termites, beetles and ants, being the most frequently detected food resource. These food resources were consumed in proportion to their relative abundance in the cerrado, suggesting that G . microtarsus is an opportunistic forager feeding primarily on insects. The generalized linear model identified sex, season and food resource as significant factors affecting the number of food items detected in faeces, as well as interactions between sex and season and season and food resource. Rate ratios calculated between sexes within seasons showed that the number of food items detected in the faeces of males was larger than that detected in the faeces of females in the warm-wet and cool-dry seasons. Rate ratios calculated between seasons within sexes showed different trends in the number of food items detected in faeces for each significant food resource. It is suggested that differences in the number of food items detected in faeces between sexes within season and between seasons within sexes are related to energetic requirements associated with reproduction.     

Food habits of Metachirus nudicaudatus (Didelphimorphia,Didelphidae) in a Brazilian Cerrado: diet composition and dietary seasonality

The diet of the brown four-eyed opossum (Metachirus nudicaudatus) at a riparian savanna in Brazil was analyzed using 22 fecal samples collected monthly from November 2009 to October 2011. A comparison was established between samples from rainy and dry seasons. Diet composition varied seasonally. Arthropods and fruits were consumed in different frequencies of occurrence, in both seasons, but small vertebrates were only found in the dry season. Diet differences among biomes (Atlantic Forest and Cerrado) were also assessed by comparing the frequency of food items in the fecal sample in our study with data from the literature. Invertebrates (100%) were the most frequent food category found, followed by fruits (59%) and vertebrates (13.6%). Small differences in the consumption of invertebrates were found among the biomes. Nevertheless, seed frequency was significantly higher in samples from the Cerrado than in those from the Atlantic Forest. In our study site, Metachirus nudicaudatus is opportunistic and has an omnivorous diet, tending to include more fruits in its natural diet than has been previously described.     

Variation of diet and food consumption of the scaldfish Arnoglossus laterna(Walbaum, 1792)

Although Arnoglossus laterna (Walbaum, 1792) is a common benthic fish in Portugal, several aspects of its feeding ecology remain incomplete. In this study, diet was examined and the food consumption estimated on inshore waters in the central coast of Portugal. The diet of the scaldfish included mainly mysids, amphipods and polychaetes, while the decapod Philocheras bispinosus Hailstone was the most frequent prey. Variation in the diet according to fish size (two size‐classes), sex and spawning/non‐spawning seasons was examined. Significant differences in diet composition were found between seasons, whereby the most frequently consumed prey in the non‐spawning season were crustaceans, while polychaetes and nemerteneans comprised most of its diet in the spawning season. The average value of the vacuity index was very similar between seasons (24.7% non‐spawning season and 25.0% spawning season). Diet overlap between size‐classes and sexes was high, but low between the two seasons. Food consumption of A. laterna was estimated based on diet characterization and evacuation rate. The estimate of the daily food consumption was 1.15 mg dry weight for the non‐spawning season and 1.67 mg dry weight for the spawning season. Significant differences in consumption rates were found between size‐classes and sexes. Larger individuals showed a higher consumption rate than smaller individuals. Females consumed more food than males in the non‐spawning season, while in the spawning season males consumed more than females.     

Does size dimorphism reduce competition between sexes? The diet of male and female pine martens at local and wider geographical scales

Sex-specific niche segregation is often used to explain sexual size dimorphism (SSD). However, whether food niche partitioning between sexes occurs as a case of sexual size dimorphism or by other mechanisms, such as behavioural dimorphism or habitat segregation, remains poorly understood. To evaluate the nature and extent of food-niche differentiation between sexes in a solitary predator I examined variation in the diet of male and female pine martensMartes martes Linnaeus, 1758 in years of high and low rodent abundance. Small mammals were the most important prey for pine martens in years of both low and high rodent abundance (occurring in more than 49% of scats). Birds, invertebrates and plant material were relatively common food items in summer diet, whereas ungulate carcasses were often consumed in autumn—winter. In general, males consumed more ungulate carcasses, plant material, amphibians and reptiles than did females, whereas females preyed more on squirrels and birds than males. There was significant seasonally dependent, between-sex variation in the occurrence of shrews, small rodents, other mammals, birds and invertebrates in marten diet. Whereas the occurrence of bank vole, birds, carcasses and plant material changed between sexes, seasons and years with various rodent abundances, both sexes consumed larger prey and had increased food niche breadth in years of low compared with high rodent abundance. Neither prey size nor food niche breadth were significantly different between males and females. The food-niche overlap between sexes was consistently lower in spring and in years of low rodent abundance. A wider geographical comparison of different marten populations showed that the diet of males and females varied significantly between locations. Females consistently preyed on squirrels and birds, whereas males fed more often on ungulate carcasses and plant material. Local and geographical comparison of male and female diets suggest that food-niche partitioning between male and female pine martens changes across different habitat and food conditions, and is not related to sexual size dimorphism, but rather to behavioural differences between sexes.     

Limitation of similarity and feeding habits in three syntopic species of newts (Triturus, Amphibia)

Differences in body size between both species and sex were observed to be close to Hutchinsonian ratios in an assemblage of three species of newts (Triturus italicus, T alpestris and T carnifex) in southern Italy According to feeding, differences in body size were not related with a strict food niche segregation The diet of larger newt included indeed all the prey eaten by smaller newts but showed new prey types Differences between species were more marked than between sexes of the same species, despite a strong sexual dimorphism in body size Larger newts foraged in a wider range of microhabitats and exhibited a greater diversity of foraging tactics Triturus italicus fed essentially on plankton and chironomids in shallow water at the banks of the pond Triturus alpestris foraged also in deeper water, particularly on lsopods in vegetation Prey fallen on the water surface constituted the main part of T carnifex diet According to recent data on newt evolution, speciation consisted in the reduction of body size Our data show that this process has been related with a specialization of the diet and of habitat use     

Behavioral Ecology of Sympatric Chimpanzees and Gorillas in Bwindi Impenetrable National Park,Uganda: Diet

Via a field study of chimpanzees (Pan troglodytes schweinfurthii) and gorillas (Gorilla gorilla beringei) in Bwindi Impenetrable National Park, Uganda, we found that their diets are seasonally similar, but diverge during lean seasons. Bwindi chimpanzees fed heavily on fruits of Ficus sp., which were largely ignored by the gorillas. Bwindi gorilla diet was overall more folivorous than chimpanzee diet, but was markedly more frugivorous than that of gorillas in the nearby Virunga Volcanoes. During 4 mo of the year Bwindi gorilla diet included more food species than that of the chimpanzees. Three factors in particular—seasonal consumption of fibrous foods by gorillas, interspecific differences in preferred fruit species, and meat consumption by chimpanzees—contributed to dietary divergence between the two species. When feeding on fruits, gorillas ate Myrianthus holstii more frequently than chimpanzees did, while chimpanzees included more figs in their annual diet. Chimpanzee diet included meat of duikers and monkeys; gorilla frequently consumed decaying wood.     

The three-dimensional macronutrient niche of an invasive generalist predator

1. Harvestmen are predators, although they suspected of being partly omnivorous by including fruits in their diet. 2. In the present study, the right-angled mixture triangle (RMT) was used to analyse the macronutritional niche of the harvestman Opilio canestrinii, an invasive species in northern Europe. The study design followed a double-test procedure in which the animals were subjected to two self-selection tests: the first one immediately after being caught in the field and the second after 1 week of ad libitum feeding during which the animals became satiated and nutritionally balanced. A comparison of results from the two tests indicates whether the animals were food limited in the field, and whether they were limited by a particular macronutrient. 3. Females were found to be food limited in the field, whereas males were not. Opilio canestrinii had a target intake of 28% lipid : 52% protein : 20% carbohydrate [i.e. with a considerable proportion of carbohydrate (sugar) in the diet]. Both sexes were non-protein limited in the field, with sugar being more limiting than lipid. 4. The results indicate considerable inclusion of plant-derived sugar in the natural diet. This conclusion was supported by a separate experiment showing enhanced performance (survival, gain in mass) in animals whose only energy supply was fresh fruit. 5. It is concluded that the harvestman is best characterized as an omnivorous predator.     

Tawny owl reproduction and offspring sex ratios under variable food conditions

Tawny owl reproduction and offspring sex ratios have been considered to depend on the abundance of small voles. We studied reproductive performance (laying date, clutch and brood size) during 1995–2003 and offspring sex ratios from 1999 to 2003 in relation to the abundance of small voles and food delivered to the nest in a tawny owl population in southern Finland. Abundance of small voles (field and bank voles) was based on trappings in the field, and estimates of food delivery was based on diet analysis of food remains in the nest boxes. In this population, reproductive output was not related to the abundance of small voles. Analysis of food delivered to the nest showed that the prey weight per offspring varied more than twofold between years and revealed that this difference was mainly related to the proportion of water voles in the diet. Only the number of water voles correlated with laying dates. Offspring sex ratios were weakly male biased (55%) but did not differ from parity. Sex ratios were not related to the abundance of small voles, and we found no evidence that parents delivered more food to nests with proportionally more offspring of the larger (female) sex. Our results underline the notion that populations may differ in their sex allocation pattern, and suggest such differences may be due to diet.     

A preliminary study of food selection by the orangutan in relation to plant quality

We observed the foraging behavior of orangutans in Central Indonesian Borneo during October, November, and December 1980, and analyzed food and nonfood items for water content, neutral detergent fiber, crude protein, available crude protein, and protein:fiber ratio and the presence of alkaloids and tannins. The diet of the orangutan during this season was unusual because it consisted predominantly of seeds and unripe, rather than ripe, fruits. Also, the major diet item, the seeds ofIrvingia malayana, had been ignored in previous years when it had fruited. In leaves, protein content was more closely associated with food choice than either neutral detergent fiber or the protein:fiber ratio. Flowers had the highest protein content and protein:fiber ratio of any food item. Tannins were found in most food items, but the presence of alkaloids was found in only one.     

Preferences of cedar waxwings in the laboratory for fruit species,colour and size: A comparison with field observations

Fruit preferences of cedar waxwings, Bombycilla cedrorum, in the laboratory were compared with preferences in the field to identify fruit characteristics that influence choice by avian dispersers. Waxwings ate 12 of 13 species of fruits offered individually in laboratory tests, but in the field ate only eight of the same 13 species. When given a choice of 10 fruit species offered simultaneously, waxwings showed a strong preference for fruits that were small or red. Preferences for fruit species in the laboratory were not correlated with morphological or nutritional characteristics of the fruits. When offered food that differed only in colour, waxwings initially preferred red over blue, yellow and green. In later tests, preference for red fruit was less marked. In tests for size preference, waxwings preferred small pieces of fruit (6 mm) over medium pieces (9 mm), and medium pieces over those that were large (12 mm). Sizes of fruits preferred in the field and in the laboratory tests were approximately 6·0–7·5 mm. Thus, waxwings can discern differences in food items and they have definite preferences. The lack of complete agreement between preferences for fruits in the field and in the laboratory suggests that factors important in the field but controlled in the laboratory (e.g. abundance, location) override preferences for certain fruits. However, some fruit characteristics, particularly size, were consistently influential in the laboratory and in the field.     

Development,survival, body weight,longevity, and reproductive potential of Oemena hirta (Coleoptera: Cerambycidae) under different rearing conditions

Oemona hirta (F.) is a New Zealand native longicorn beetle, whose larvae bore into the wood of branches and stems of living trees and vines, causing serious damage. To explore effective methods for maintaining laboratory colonies and biology of immature stages of this species we evaluated four laboratory rearing methods with both natural and artificial diets and compared biological parameters of laboratory colonies with those of field-collected insects. On an artificial diet, approximately 40% of neonate and 70% of autumn- and 11% of winter-collected larvae reached adulthood. Neonate larvae could not complete their development in cut poplar (Populus nigra variety italica Koehne) twigs; however, when twigs were standing in water >46% of neonate larvae survived to adulthood. Mean larval development time ranged from approximately150 to almost 300 d, depending on rearing methods. Mean pupal stage ranged from 15 to 19 d. Adult females were significantly heavier than males. Although adult females from field-collected twigs and reared on the artificial diet had similar mean body weight, which was significantly greater than that of other rearing colonies, the potential fecundity was the highest in the former (83.1 +/- 29.4 eggs) and the lowest in the latter (33.5 +/- 9.1 eggs). In all laboratory-reared and field-collected insects, the total number of eggs produced (eggs laid + eggs that remained in dead female body) by females was positively correlated with their body weight. When larvae fed on the artificial diet, there was no positive correlation between the number of eggs successfully laid and female body weight. However, when larvae lived on natural food, a positive correlation was found. In laboratory colonies, mean longevity of females (36 to 52 d) was slightly greater than that of males (30 to 50 d) without significant difference between sexes but in the adults from field-collected twigs, males (52 d) lived significantly longer than females (33 d). In terms of time, labor, and the number of resulting adults, collecting larvae in the field in autumn and then transferring them onto artificial diet is the most effective method for maintaining a laboratory colony.     

Molecular detection of invertebrate prey in vertebrate diets: trophic ecology of Caribbean island lizards

Understanding community assembly and population dynamics frequently requires detailed knowledge of food web structure. For many consumers, obtaining precise information about diet composition has traditionally required sacrificing animals or other highly invasive procedures, generating tension between maintaining intact study populations and knowing what they eat. We developed 16S mitochondrial DNA sequencing methods to identify arthropods in the diets of generalist vertebrate predators without requiring a blocking primer. We demonstrate the utility of these methods for a common Caribbean lizard that has been intensively studied in the context of small island food webs: Anolis sagrei (a semi‐arboreal ‘trunk‐ground’ anole ecomorph). Novel PCR primers were identified in silico and tested in vitro. Illumina sequencing successfully characterized the arthropod component of 168 faecal DNA samples collected during three field trips spanning 12 months, revealing 217 molecular operational taxonomic units (mOTUs) from at least nine arthropod orders (including Araneae, Blattodea, Coleoptera, Hemiptera, Hymenoptera, Isoptera, Lepidoptera and Orthoptera). Three mOTUs (one beetle, one cockroach and one ant) were particularly frequent, occurring in ≥50% of samples, but the majority of mOTUs were infrequent (180, or 83%, occurred in ≤5% of samples). Species accumulation curves showed that dietary richness and composition were similar between size‐dimorphic sexes; however, female lizards had greater per‐sample dietary richness than males. Overall diet composition (but not richness) was significantly different across seasons, and we found more pronounced interindividual variation in December than in May. These methods will be generally useful in characterizing the diets of diverse insectivorous vertebrates.     

Variations in diet composition of sympatric Trachypithecus francoisi and Macaca assamensis in the limestone habitats of Nonggang,China

Comparative studies of sympatric species are essential for understanding behavioral and ecological adaptation as well as the mechanisms that can reduce resource competition to allow coexistence.Fran?ois' langurs(Trachypithecus francoisi) and Assamese macaques(Macaca assamensis) are sympatric primate species found in the limestone seasonal rainforests of Nonggang Nature Reserve,southwestern Guangxi, China. To explore their different adaptation strategies, we collected data on diet using scan sampling at 15-min intervals. Our results revealed that Fran?ois' langurs showed a more flexible diet composition than Assamese macaques.Fran?ois' langurs increased dietary diversity and mature leaf consumption in response to seasonal scarcity of preferred young leaves and fruits, whereas Assamese macaques relied heavily on young bamboo leaves(Indocalamus calcicolus) in most months.These variations reflect the differences in digestive physiology, morphology, and the temporal and spatial distribution of food resources.     

Seasonal and between-sex differences in the diet of Black Grouse Tetrao tetrix

Capsule Dietary differences between sexes and seasons reflected diversity in plant availability and habitat preferences.

Aims To analyse Black Grouse diet through the identification of plant and invertebrate material found in the crop.

Methods Crops were dissected and the content categorized into plant species and components (i.e. leaf, stem, flower, fruit and seed). Multivariate analysis was used to test for seasonal, sex- or location-related differences in the diet.

Results Plant fragments accounted for 98% of the diet and included 53 plant species or taxa. Invertebrates made up the remaining 2%. Diet varied significantly between seasons and sexes. Both sexes ate more ericaceous shrubs in autumn and winter, with females eating more than males. The plant parts eaten varied seasonally. In summer, fruits, flowers and seeds were favoured over leaves, which dominated in winter.

Conclusions With few trees, birds were reliant upon Heather in autumn and winter.     

Assessing the Occurrence of Sexual Segregation in Spider Monkeys (Ateles geoffroyi yucatanensis), Its Mechanisms and Function

Sexual segregation is a recognized dimension of the socioecology of many vertebrates, but it has not been systematically examined in primates. We investigated temporal patterns of sexual segregation in spider monkeys (Ateles geoffroyi yucatanensis) using a test that distinguishes sexual segregation from aggregation and random association between the sexes. We further investigated how sexual segregation varies over time as a function of food availability, and then tested other possible factors that might be causally linked to sexual segregation in spider monkeys. We predicted that male philopatry and cooperative territorial defence leads to sexual dimorphism in behavior, which in turn creates different optimal energetic requirements for males and females as reflected in differing activity budgets and diet. We investigated sexual segregation in a group of 33–35 spider monkeys at Runaway Creek Nature Reserve in Belize over 23 mo in 2008–2009. We used the sex compositions of subgroups recorded in scan samples to test the occurrence of sexual segregation at monthly and biweekly time scales.We found that males and females were significantly segregated in 15 out of the 23 mo of the study, and that periods of nonsegregation coincided with months of low food availability. The sexes differed significantly in activity and diet; males spent more time traveling, and less time resting and feeding than females, and they had a higher proportion of ripe fruits in their diets than did females. We propose that sexual segregation in spider monkeys is primarily a form of social segregation that results from males and females pursuing optimal dietary and behavioral strategies to satisfy sex-specific energetic demands. We further suggest that sexual segregation represents an important constraint on fission–fusion dynamics that should be considered when assessing the potential for variability in subgroup composition.     

Which proximate factor determines sexual size dimorphism in tiger snakes?

Diverse interactions between factors that influence body size complicate the identification of the primary determinants of sexual size dimorphism. Using data from a long‐term field study (1997–2009), we examined the contributions of the main proximate factors potentially influencing sexual size dimorphism from birth to adulthood in tiger snakes (Notechis scutatus). Data on body size, body mass and body condition of neonates, juveniles and adults were obtained by mark–recapture. Frequent recaptures allowed us to monitor reproductive status, diet and food intake, and to estimate survival and growth rates in age and sex classes. Additional data from females held briefly in captivity enabled us to assess reproductive output and the body mass lost at parturition (proxies for reproductive effort). From birth to maturity, individuals of both sexes experienced similar growth and mortality rates. We found no difference in diet, feeding and survival rates between the sexes, nor between juveniles and adults. On maturity, despite comparable diet and food intake by both sexes, the high energy requirements of vitellogenesis and gestation were responsible for a depletion of body reserves and probably resulted in a marked decrease in growth rates. Males were largely exempt from such costs of reproduction, and so could grow faster than females and attain larger body sizes. The absence of niche divergence between the sexes (uniformity of habitat, lack of predators) suggests that the impact of differential energetic investment for reproduction on growth rate is probably the main proximate factor influencing sexual size dimorphism in this species. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 668–680.     

Food habits and space-use of red pandas <Emphasis Type="Italic">Ailurus fulgens</Emphasis> in the Fengtongzhai Nature Reserve,China: food effects and behavioural responses

The red panda Ailurus fulgens Cuvier, 1825 is endemic to the Himalayan-Hengduan Mountains. Many of its ecological traits are still poorly understood although important from a scientific standpoint as a highly specialized bamboo feeder. To understand its food habits and space-use, we carried out a field study from April 2002 to May 2003 in Fengtongzhai Nature Reserve, China. The results showed that red pandas almost exclusively fed on Bashania faberi, but not on Yushania brevipaniculata in the study area. Besides differences in nutritional quality, accessibility to food items is perhaps another factor affecting their utilization of bamboo species. B. faberi leaves are the highest in crude protein, being the primary year-round food source; new shoots and fruits of Sorbus, and Rubus are nutritious and digestible, seasonally occurred in their diet. Red pandas seemed to select most nutritious food items to feed on. The home range of red pandas averaged 2.2 km2 across individuals (2.6 km² for the males and 1.7 km² for the females), and overlapped extensively (ranging 18.9–78.1%). No significant difference was found in monthly percentage of total home range used among seasons and sexes. The daily movement distance was 455 m (447 m for the females and 463 m for the males), significantly affected by sexes and months. Our findings indicated that food resource significantly influenced behaviour of red pandas in foraging and space use, and their behavioural responses are adaptive being helpful to energy intake and maintenance.     

Seasonal and inter‐island variation in the foraging strategy of the critically endangered Red‐headed Wood Pigeon Columba janthina nitens in disturbed island habitats derived from high‐throughput sequencing

We studied the feeding ecology of the critically endangered Red‐headed Wood Pigeon Columba janthina nitens, a subspecies endemic to a very remote and highly disturbed oceanic island chain, the Ogasawara Islands. An analysis based on high‐throughput sequencing (HTS) was undertaken on 627 faecal samples collected over 2 years from two island habitats, and food availability and the nutrient composition of the major fruits were also estimated. The HTS diet analysis detected 122 food plant taxa and showed clear seasonal and inter‐island variation in the diet of the Pigeons. The results indicated a preference for lipid‐rich fruits, but the diet changed according to the availability of food resources, perhaps reflecting the foraging strategy of the Pigeons in isolated island habitats with poor food resources. Pigeons also frequently consumed introduced plants at certain times of year, perhaps compensating for the lack of preferred native food resources. However, the degree of dependence on introduced plants appeared to differ between the two island habitats, so the different impacts of introduced plant eradication on the foraging conditions for the Pigeons on each island should be considered. HTS diet analysis combined with field data may be useful for monitoring the foraging conditions of endangered species and may also inform an appropriate conservation strategy in oceanic island ecosystems with complicated food webs that include both native and introduced species.