Social and demographic influences on the feeding ecology of giraffe in the Luangwa Valley,Zambia: 1973–2014

Body size influences metabolic rate, which impacts feeding ecology. Body mass differs by sex in size‐dimorphic species, such as giraffes, and also by age. Giraffes reside in a fission–fusion social system, which influences feeding ecology due to frequent changes in group size and composition. We analysed 40 years of feeding records collected from a population of Thornicroft's giraffes (Giraffa camelopardalis thornicrofti) living in the Luangwa Valley, Zambia. We examined the influence of herd composition, age and sex on diet. Solitary males and herd males did not differ in diet. Dietary diversity was comparable for females and males, with sex differences in plant species eaten present during the dry season. Age differences in feeding ecology were pronounced, with juveniles often feeding on bushes and smaller trees, while adults tended to feed upon taller trees. Both sexes have evolved foraging strategies that maximize nutrient and energy intake commensurate with their reproductive strategies, with male metabolic requirements sometimes greater, and sometimes less, than that of females. We propose that females are not exchanging foraging efficiency with reproductive tactics by feeding on smaller trees in the open, but are increasing prospects for their calves to survive when confronted with interspecific competition by browsers.     

Wet and dry season diets of the Endangered Lemur catta (ring‐tailed lemur) in two mountainous rocky outcrop forest fragments in south‐central Madagascar

Lemur catta's ability to consume a wide variety of plant foods is a key to this species' survival in a time of ecological crisis across its geographic range. We examined seasonal diet variability of L. catta groups inhabiting two rocky outcrop fragments in south‐central Madagascar: Anja Reserve and Tsaranoro Valley forest. Leaves and fruit of Melia azedarach were a keystone resource for Anja lemurs in wet and dry seasons. At Tsaranoro, L. catta relied on M. azedarach and Ficus spp. in dry season, but during the wet season, neither was a dominant food resource. Top food species at both sites differed markedly from those consumed by L. catta in other habitats. At Tsaranoro, a greater proportion of lemurs engaged in feeding during the dry season compared with wet season. We attribute this to resource scarcity during dry season, when greater feeding effort is needed to maintain energy requirements. Because M. azedarach is ubiquitous throughout Anja Reserve, producing fruit and leaves year‐round, Anja lemurs can meet energy requirements with little seasonal adjustment in feeding activity. L. catta's IUCN status has been upgraded to Endangered, thus, greater insight into its diet flexibility and ability to survive on introduced plant species, can inform conservation plans in remaining wild habitats and ex situ programs.     

Mitochondrial DNA analyses show that Zambia's South Luangwa Valley giraffe (Giraffa camelopardalis thornicrofti) are genetically isolated

Thornicroft's giraffe, Giraffa camelopardalis thornicrofti, is a geographically isolated subspecies of giraffe found only in north‐east Zambia. The population only occurs in Zambia's South Luangwa Valley, an area which interestingly places it between the current distribution of Masai (G. c. tippelskirchi) giraffe to the north, and the Angolan (G. c. angolensis) and South African (G. c. giraffa) giraffe in the south‐west and south, respectively. Specific studies have been undertaken on the ecology of this subspecies, but their population genetics remains unknown. We studied 34 individuals from the South Luangwa National Park and adjacent Lupande Game Management Area and seven individuals from northern Botswana. The complete cytochrome b and control region sequences of the mitochondrial genome were sequenced and analysed together with database data by maximum likelihood tree reconstruction and maximum parsimony network analyses. The giraffe from Zambia's South Luangwa Valley are most closely related to the subspecies G. c. tippelskirchi and part of their radiation. However, they form a unique population that would benefit from increased research and conservation management.     

Hierarchical foraging by giraffe in a heterogeneous savannah,Tanzania

Understanding foraging decisions made by wildlife at different spatio‐temporal scales is important for wildlife management and conservation. We tested whether foraging decisions by Masai giraffe (Giraffa camelopardalis tippelskirchi Matschie) differed with scales; habitat, patch and tree in a heterogeneous savannah. We collected data from Arusha National Park, Tanzania, in March–May and August–October 2013. Visual observations were used to collect data on foraging behaviour. Measurements of tree height and stem height and scoring of accumulated browsing were made in 133 patches around trees where the giraffes had been seen browsing, and in a corresponding number of available patches. Giraffes selected Acacia shrub and Dodonea shrub habitats but not for water availability or predator avoidance. For patch use, giraffe selected high quantity of Acacia xanthophloea and Dodonea viscosa. Between plant species, A. xanthophloea was the most preferred and within plant species, tree quality was enhanced by tree height and high score of accumulated browsing. Generally, giraffes selected for A. xanthophloea at all scales.     

Species‐specific foraging strategies and segregation mechanisms of sympatric Antarctic fulmarine petrels throughout the annual cycle

Determining the year‐round distribution and behaviour of birds is necessary for a better understanding of their ecology and foraging strategies. Petrels form an important component of the high‐latitude seabird assemblages in terms of species and individuals. The distribution and foraging ecology of three sympatric fulmarine petrels (Southern Fulmar Fulmarus glacialoides, Cape Petrel Daption capense and Snow Petrel Pagodroma nivea) were studied at Adélie Land, East Antarctica, by combining information from miniaturized saltwater immersion geolocators and stable isotopes from feathers. During the breeding season at a large spatial scale (c. 200 km), the three species overlapped in their foraging areas located in the vicinity of the colonies but were segregated by their diet and trophic level, as indicated by the different chick δ¹⁵N values that increased in the order Cape Petrel < Southern Fulmar < Snow Petrel. During the non‐breeding season, the three fulmarines showed species‐specific migration strategies along a wide latitudinal gradient. Snow Petrels largely remained in ice‐associated Antarctic waters, Southern Fulmars targeted primarily the sub‐Antarctic zone and Cape Petrels migrated further north. Overall, birds spent less time in flight during the non‐breeding period than during the breeding season, with the highest percentage of time spent sitting on the water occurring during the breeding season and at the beginning of the non‐breeding period before migration. This activity pattern, together with the δ¹³C values of most feathers, strongly suggests that moult of the three fulmarine petrels occurred at that time in the very productive high Antarctic waters, where birds fed on a combination of crustaceans and fish. The study highlights different segregating mechanisms that allow the coexistence of closely related species, specifically, prey partitioning during the breeding season and spatial segregation at sea during the non‐breeding season.     

Dietary overlap and seasonality in three species of mormoopid bats from a tropical dry forest

Competing hypotheses explaining species’ use of resources have been advanced. Resource limitations in habitat and/or food are factors that affect assemblages of species. These limitations could drive the evolution of morphological and/or behavioural specialization, permitting the coexistence of closely related species through resource partitioning and niche differentiation. Alternatively, when resources are unlimited, fluctuations in resources availability will cause concomitant shifts in resource use regardless of species identity. Here, we used next‐generation sequencing to test these hypotheses and characterize the diversity, overlap and seasonal variation in the diet of three species of insectivorous bats of the genus Pteronotus. We identified 465 prey (MOTUs) in the guano of 192 individuals. Lepidoptera and Diptera represented the most consumed insect orders. Diet of bats exhibited a moderate level of overlap, with the highest value between Pteronotus parnellii and Pteronotus personatus in the wet season. We found higher dietary overlap between species during the same seasons than within any single species across seasons. This suggests that diets of the three species are driven more by prey availability than by any particular predator‐specific characteristic. P. davyi and P. personatus increased their dietary breadth during the dry season, whereas P. parnellii diet was broader and had the highest effective number of prey species in all seasons. This supports the existence of dietary flexibility in generalist bats and dietary niche overlapping among groups of closely related species in highly seasonal ecosystems. Moreover, the abundance and availability of insect prey may drive the diet of insectivores.     

Diets of the African Civet Civettictis civetta (Schreber, 1778) in selected coffee forest habitat,south‐western Ethiopia

Feeding ecology of the African civet, Civettictis civetta (Schreber, 1778), was studied in selected coffee forest habitat for two seasons between December 2012 and October 2013. The study was conducted in Limu Seka district, south‐west Ethiopia. Faecal analysis was employed to assess the diet spectrum, seasonal abundance and relative importance of food items. Over 55 different food items were identified from the analysis of 387 scat samples collected during the dry and wet seasons. In the coffee forest habitat, African civets showed omnivorous feeding habit with plant‐to‐animal diet biomass ratio of 1 : 1.36. A slight diet specialization was observed during the wet season (BA = 0.46) favouring animal prey. However, during the dry season, they showed generalist feeding habit (BA = 0.87) with more plant biomass in their diet. With over 64% occurrence and 14.4% biomass, coffee berries significantly contributed to the civets dry season plant diets. The excreted coffee beans, after civets ingested ripe coffee berries, are the tastiest product used by farmers for consumption and market. Seasonal collection of civet coffee from coffee forest floor economically supports the farmers while increasing the importance of civets in the habitat and hence contributing for its conservation.     

Feeding habits of the speckled guitarfish Rhinobatos glaucostigma (Elasmobranchii: Rhinobatidae)

The stomachs of 464 speckled guitarfish Rhinobatos glaucostigma were sampled from the south‐eastern Gulf of California (GC) to determine diet composition. Numerical indices and prey‐specific index of relative importance (%I_PSIR) were used to determine the feeding strategy of the species. An analysis of similarity (ANOSIM) was used to determine differences in diet with respect to sex, season (dry or rainy) and maturity stages (immature or mature). The diversity and niche breadth (by sex, season and maturity) and a general trophic level were determined. The overall diet was dominated by shrimps (%I_PSIR = 43·47), amphipods (%I_PSIR = 18·89) and crabs (%I_PSIR = 18·07). ANOSIM demonstrated differences in the diet by maturity and season, but not by sex. Rainy and dry season diets were dominated by shrimps and amphipods, respectively. Immature specimens fed mainly on amphipods, whereas mature fish preferred shrimps and crabs. Rhinobatos glaucostigma showed a narrow niche breadth with an intermediary trophic level (T_L = 3·72) and can be considered as a secondary consumer in the soft‐bottom demersal community of the south‐east GC. Understanding the feeding habits and trophic level of R. glaucostigma is vital to help identify the segments of the population vulnerable to overfishing by artisanal and industrial fisheries, and to aid in conservation and management of this elasmobranch.     

Seasonal variation in food availability and relative importance of dietary items in the Gambian epauletted fruit bat (Epomophorus gambianus)

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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.     

Discrimination of grasshopper (Orthoptera: Acrididae) diet and niche overlap using next‐generation sequencing of gut contents

Species of grasshopper have been divided into three diet classifications based on mandible morphology: forbivorous (specialist on forbs), graminivorous (specialist on grasses), and mixed feeding (broad‐scale generalists). For example, Melanoplus bivittatus and Dissosteira carolina are presumed to be broad‐scale generalists, Chortophaga viridifasciata is a specialist on grasses, and Melanoplus femurrubrum is a specialist on forbs. These classifications, however, have not been verified in the wild. Multiple specimens of these four species were collected, and diet analysis was performed using DNA metabarcoding of the gut contents. The rbcLa gene region was amplified and sequenced using Illumina MiSeq sequencing. Levins' measure and the Shannon–Wiener measure of niche breadth were calculated using family‐level identifications and Morisita's measure of niche overlap was calculated using operational taxonomic units (OTUs). Gut contents confirm both D. carolina and M. bivittatus as generalists and C. viridifasciata as a specialist on grasses. For M. femurrubrum, a high niche breadth was observed and species of grasses were identified in the gut as well as forbs. Niche overlap values did not follow predicted patterns, however, the low values suggest low competition between these species.     

Feeding habit and food partitioning in a stream fish community of Western Ghats,India

The diet and feeding ecology of 12 cyprinid fishes were studied in the Thalayanai stream of southern India. The diet of Barbodes sarana, Danio aequipinnatus and Rasbora daniconius were specialized in terrestrial and aquatic insects. Hypselobarbus dobsoni and Garra mullya consumed vegetative matter and unicellular algae as their most important dietary component, respectively. Hypselobarbus curmuca and Hypselobarbus dubius specialized in crustaceans and molluscs. Puntius amphibious, Puntius bimaculatus, Puntius dorsalis, Puntius filamentosus and Puntius tambraparniei fed on variety of food resources (aquatic insects, algae and detritus) and their feeding intake varied between wet to dry seasons. Levin’s index of diet breadth showed the broadest diets for P. dorsalis in both seasons, whereas G. mullya had a narrow feeding spectrum in both seasons. Among co-existing species, diet overlaps were substantially low during both wet and dry season. These results suggest that feeding specialization and low diet overlap reduce inter-specific competition, allowing co-existence of these cyprinid fishes in this highly structured stream assemblage.     

Foraging Patterns of Sympatric Hornbills during the Nonbreeding Season in Arunachal Pradesh, Northeast India

The diet and food resource partitioning of three sympatric hornbills (Great Hornbill Buceros bicornis, Wreathed Hornbill Aceros undulatus, and Oriental Pied Hornbill Anthracoceros albirostris) during the nonbreeding season were studied relative to fruit availability in a foothill semievergreen forest of Arunachal Pradesh, northeast India. Hornbills fed on fruits of 49 plant species that comprised over 95 percent of their diet. Hornbill species partitioned food resources by varying the relative contribution of figs and non‐fig fruits in the diet. Similarity in non‐fig fruit diet was low. Ten species contributed to over 90 percent of the non‐fig diet. The availability of non‐fig fruits was much lower in the nonbreeding season than in the breeding season; however, despite lower fruit availability during the nonbreeding season, hornbills had a wide diet breadth and indirect evidence suggests that Wreathed Hornbills foraged widely in this season for fruit. Great Hornbills relied more on figs that were available year‐round. The diverse diet breadth that included rare and patchy fruit resources underscores the importance of conserving large forest tracts for hornbills.     

Extreme trophic segregation between sympatric fish species: the case of small sized body <Emphasis Type="Italic">Aphyocharax</Emphasis> in the Brazilian Pantanal

Interspecific differences in diet of the two sympatric fish species of the subfamily Aphyocharacinae were investigated. Both were sampled monthly (March 2000 to February 2001) during the rainy and dry season in North region of the Pantanal. Aphyocharax anisitsi was an invertivorous/insectivorous species, feeding mainly on aquatic insects during the rainy season and terrestrial insects in the dry season. Aphyocharax dentatus remained a piscivorous regardless of the hydrological season. Low feeding-overlap values, 0.079 in the dry season and no overlap in the rainy season were observed. The diet-breadth values for each species were low (less than 0.1) except for A. anisitsi in the lake during the dry season (0.4). A detrended correspondence analysis (DCA) summarized the diet similarity patterns: in DCA 1, it was observed that fish prey grouped A. dentatus in the lower scores, and insect prey grouped A. anisitsi in the higher scores. A two-way ANOVA showed that the diets differed significantly between the species. Our results show a clear trophic segregation between these species, supporting the niche concept that presumes that differences between sympatric species must exist, to allow their coexistence. Besides extreme segregation, it was clear that A. dentatus is a piscivorous fish, despite its low size (until 72 mm). This finding counteracts the current view about food web structure that poses a direct and positive relationship between body size and trophic position.     

Feeding habits of the Chinese minnow Rhynchocypris oxycephalus (Sauvage & Dabry de Thiersant, 1874) from the upper Yangtze River,China

The feeding habits of the Chinese minnow, Rhynchocypris oxycephalus (Sauvage & Dabry de Thiersant, 1874), were investigated in the Laohegou River, a tributary of the upper Yangtze River, China. A total of 412 specimens were collected by electrofishing in four seasons of 2012, 7 days per season. Diet composition was analyzed according to season, size classes and sex. The Chinese minnow is a generalist omnivorous species, whose diet consists of aquatic insects and plants. Besides algae, Hemiptera, Ephemerida, Trichoptera, Odonata, Diptera were the most dominat food items. The Shannon‐Wiener index on diet items showed no significant differences between major food items (p > .05) or between seasons, size classes and sexes. In contrast, fullness index and dietary breadth analysis both indicated significant diet changes between seasons. Consequently, this study provides information on feeding habits of R. oxycephalus, which may be useful in cultivation trials.     

Seasonal and individual predictors of diet in a free‐ranging population of mandrills

Deciphering the dietary habits of a species is central to understanding its ecology, resource requirements, and the evolution of its life history traits. Detailed information on how primates use their environment to address their nutritional needs is available for many primate species. Such basic, but necessary data are, however, fragmented for secretive primates, especially regarding direct behavioral observations of individuals. In this study, we evaluated the impact of seasonality and demographic characteristics on diet and feeding habits in the only free‐ranging population of habituated mandrills (Mandrillus sphinx), a forest‐dwelling species inhabiting the dense humid forests of Central Africa. We collected fine‐grained quantitative data on feeding behavior of 57 individually‐recognized animals of both sexes and different age classes during a 17‐month period. We identified most consumed plant species and determined their abundance in the habitat of the studied mandrills. We showed that diet in this species was extremely diverse and included approximately 150 different plant species, but also mushrooms, invertebrates, and vertebrates. This omnivorous and highly diverse diet presented, however, a clear frugivorous tendency. While we identified three food items largely consumed throughout the year, we also found a strong seasonal signature on diet that was partly, but not only, related to food availability. Age and sex also influenced feeding habits with some feeding specializations according to the individual categories considered and their associated nutritional needs. Our quantitative data provide a basis for future studies examining the nutritional and mineral content of food items, which will further elucidate important aspects of the ecology of this little studied forest primate.     

Reproductive phenology of two Mimusops species in relation to climate,tree diameter and canopy position in Benin (West Africa)

Assessing species phenology provides useful understanding about their autecology, to contribute to management strategies. We monitored reproductive phenology of Mimusops andongensis and Mimusops kummel, and its relationship with climate, tree diameter and canopy position. We sampled trees in six diameter classes and noted their canopy position. For both species flowering began in the dry season through to the rainy season, but peaked in the dry season, whilst fruiting occurred in the rainy season and peaked during the most humid period. Flowering was positively correlated with temperature. Conversely, fruiting was negatively correlated with temperature and positively with rainfall, only in the Guineo‐Sudanian zone. For M. andongensis, flowering and fruiting prevalences were positively linked to stem diameter, while only flowering was significantly related to canopy position. For M. kummel, the relationship with stem diameter was significant for flowering prevalence only and in the Guineo‐Sudanian zone. Results suggest that phylogenetic membership is an important factor restricting Mimusops species phenology. Flowering and fruiting of both species are influenced by climate, and consequently climate change might shift their phenological patterns. Long‐term investigations, considering flowering and fruiting abortion, will help to better understand the species phenology and perhaps predict demographic dynamics.     

Dietary specialization and the effects of plant species on potential multitrophic interactions of three species of nymphaline caterpillars

The diet breadth of insect herbivores influences their response to variation in plant quality, and these bitrophic interactions have implications for the higher‐level trophic interactions between herbivores and their natural enemies. In this comparative study, we examined the role of host plant species and plant secondary chemistry on the potential interactions between three species of nymphaline caterpillars and their natural enemies. The caterpillar species (all Lepidoptera: Nymphalidae) varied in their degree of specialization: the buckeye, Junonia coenia Hübner, is a specialist on plants that contain iridoid glycosides (IGs); the white peacock, Anartia jatrophae L., feeds on plants in five families, some of which contain IGs and some of which do not; and the painted lady, Vanessa cardui L., is a generalist, feeding on plants in at least 15 families. Each species was reared on leaves of an introduced host plant, Plantago lanceolata L. (Plantaginaceae), which produces two IGs, aucubin and catalpol, and on another plant species that is a common host plant. These alternate host plants were Plantago major L. (Plantaginaceae) for J. coenia, Bacopa monnieri (L.) Pennell (Plantaginaceae) for A. jatrophae, and Malva parviflora L. (Malvaceae) for V. cardui. We examined growth, sequestration, and immune response of these caterpillars on the different host plant species. Junonia coenia developed more rapidly and sequestered higher IG concentrations when reared on P. lanceolata, whereas both other species grew more slowly on P. lanceolata. Host plant did not influence immune response of J. coenia or A. jatrophae, whereas V. cardui immune response was weaker when reared on P. lanceolata. Junonia coenia was most efficient at IG sequestration and A. jatrophae was least efficient, when all three species were reared on P. lanceolata. These results indicate that diet breadth may play an important role in structuring tritrophic interactions, and this role should be further explored.     

Influence of flood pulses on diet composition and trophic relationships among piscivorous fish in the upper Paraná River floodplain

The aim of this article is to evaluate whether alterations in flood pulses differentially affect diet composition, feeding niche breadth, and diet overlap of piscivorous fish. Species examined were Acestrorhynchus lacustris, Hoplias aff malabaricus, Plagioscion squamosissimus, Rhaphiodon vulpinus, and Salminus brasiliensis. These species were collected with gillnets (different mesh sizes) in the upper Paraná River floodplain, during four distinct flood events (four periods; A = 1992/1993; B = 2000; C = 2001; and D = 2002). The volumetric method was chosen to express diet results. Feeding niche breadth was calculated using Levins measure, and diet overlap was evaluated by the Pianka’s Index. Flooding was more intense and lasted longer in the first period (1992/1993—A). Diet composition of the studied species was broad (47 total items consumed). For period A, Prochilodus lineatus was the main item taken by four out of five species. In the other periods, there were relevant alterations in diet, since P. lineatus was not recorded in any stomach of the five species; rather, it was replaced by the shrimp, Macrobrachium amazonicum. Diet overlap was low in all periods. The greatest overlap was obtained in period C for P. squamosissimus and R. vulpinus, due to high consumption of shrimps. There were no significant differences in niche breadth among species. However, the species presented distinct variation patterns in niche breadth. For example, H. aff. malabaricus showed a tendency toward increasing niche over the period, but the other species presented larger niches only during period A. Therefore, it can be concluded that the intensity and duration of the flood pulse influences: (i) the diet composition of piscivores; (ii) the breadth of their niches; and (iii) feeding overlap among species. Handling editor: J. Cambray     

Not all ants are equal: obligate acacia ants provide different levels of protection against mega‐herbivores

In obligate ant–plant mutualisms, the asymmetric engagement of a single plant species with multiple ant species provides the opportunity for partners to vary in their behaviour. Variation in behaviour has implications for the interactions with third‐party species such as herbivores. This study assessed the effect of obligate ant‐mutualists (Crematogaster mimosae, Crematogaster nigriceps and Tetraponera penzigi) inhabiting the African ant‐acacia (Acacia drepanolobium) on three mega‐herbivore browsers: the Maasai giraffe (Giraffa camelopardalis tippelskirchi), the reticulated giraffe (Giraffa c. reticulata) and the black rhino (Diceros bicornis). Giraffes are abundant and wide‐ranging herbivores of the acacias, whereas black rhinos are localized and perennial herbivores of the acacias. Multiyear field studies comparing the ants’ aggressive behaviour and browsing by mega‐herbivores suggested differences between the tending abilities of the primary ant species inhabiting A. drepanolobium. Trees occupied by the aggressive ant species C. mimosae had significantly less browsing by giraffes and black rhino than trees occupied by other ant species. The results of this study provide evidence that ant‐mutualists on African acacias can serve as deterrents to mega‐herbivores and that different ant species vary in their tending abilities.