The importance of seed dispersal in the Alexandria Coastal Dunefield, South Africa

The endozoochorous dispersal of seeds by mammals and birds between distinct vegetation communities was assessed to determine the importance of these processes in coastal dune field management. Isolated pockets of thicket vegetation (bush-pockets) within a large coastal dune field provided the opportunity to study vertebrate seed dispersal and its contribution to their origin and maintenance. Mammalian and avian faeces were collected for the quantification of seeds dispersed via endozoochory. Birds and mammals showed considerable overlap, dispersing intact seeds of 17 and 29 plant species, respectively, but mammals dispersed a greater diversity and size range than birds. Extrapolation of mammalian faecal data indicates an annual input of 23 million intact seeds to the dune field. Significantly more seeds are deposited by mammals and birds in the bush-pockets than on open sand, and birds deposited greater numbers of seeds nearer the seed source. Zoochory appears to be critical for the maintenance of the bush-pocket habitats through the dispersal of climax woody plant species into the dune field. Directional dispersal by birds and mammals to the bush-pockets is considered to be responsible for the maintenance and possible origin of these bush-pockets. The high number of exotic plant propagules dispersed by both avian and mammalian zoochory highlights the importance of management of the Alexandria Coastal Dunefield (ACD) beyond the reserve boundaries. In a dynamic system such as the ACD which is within a declared nature reserve, the continued existence of the bush-pockets may depend on the maintenance, beyond the reserve boundaries, of a reservoir of not only plant material but vertebrate dispersers as well.     

Flexible seed selection by individual harvester ants, Pogonomyrmex occidentalis

Summary Individual seed harvester ants (Pogonomyrmex) have been shown to specialize on specific seed types. We examined possible mechanisms for seed specialization and tested whether fidelity to food type limits the foraging decisions of individual western harvester ants, Pogonomyrmex occidentalis. The seed selection regimes of individually marked ants foraging at piles of two seed types were described and related to differences in seed quality and colonial dietary history. Individual foraging choices were affected by multiple factors, including seed caloric rewards, the previous seed selected, and the dietary history of the colony. Individual seed choices generally converged on the most energetically profitable species, suggesting that foragers exhibit labile preference. However, for a portion of the foragers, seed specialization was also partially due to constancy, defined as a tendency to select seed species that were previously collected. When colonies were presented with one seed type for 1 h and then were offered a mix of that seed and a novel seed type, individuals showed a strong preference for the novel seeds. Such rapid changes in seed preference argue strongly that individual P. occidentalis ants are highly flexible in seed choice and that resource assessment by these ants is more complex than simple maximization of net energetic return.Offprint requests to: J.H. Fewell at the current address     

Evaluating day–night changes in shallow Mediterranean rocky reef fish assemblages by visual census

Ecological information on coastal fish distribution patterns and habitat use can be greatly improved by nocturnal samplings and observations. To this purpose, the structure of a Mediterranean fish assemblage inhabiting the shallow rocky littoral of Linosa Island (Sicily Strait, Italy) was examined by using visual census to detect possible diel variations in species composition and abundance. Day–night fish distribution patterns were investigated by multivariate and univariate analyses. Overall, 42 fish taxa belonging to 19 families were recorded: 35 during the day and 24 during the night. Seventeen species were common to both diurnal and nocturnal assemblages. Within the diurnal assemblage, Chromis chromis was the most represented species (37.2%), followed by Thalassoma pavo (23.2%) and Sparisoma cretense (10.8%). Within the nocturnal assemblage, the most abundant taxon was Atherina spp. (33.9%), followed by Apogon imberbis (26.4%) and Boops boops (11.5%). Our results indicated wide variation in the abundance and species composition during the day and during the night. Multi Dimensional Scaling plot showed a clear-cut separation between the two assemblages and analysis of similarities found significant differences as well. SIMPER analysis revealed that ten species individually contributed by more than 2.5% to the dissimilarity between diurnal and nocturnal assemblages, T. pavo, C. chromis and S. cretense being the first three species in order of decreasing percentage. ANOVA performed on species richness and fish abundance detected significant differences between diurnal and nocturnal assemblages, the latter showing far lower average values for both variables.     

Separating selection by diurnal and nocturnal pollinators on floral display and spur length in Gymnadenia conopsea

Most plants attract multiple flower visitors that may vary widely in their effectiveness as pollinators. Floral evolution is expected to reflect interactions with the most important pollinators, but few studies have quantified the contribution of different pollinators to current selection on floral traits. To compare selection mediated by diurnal and nocturnal pollinators on floral display and spur length in the rewarding orchid Gymnadenia conopsea, we manipulated the environment by conducting supplemental hand-pollinations and selective pollinator exclusions in two populations in central Norway. In both populations, the exclusion of diurnal pollinators significantly reduced seed production compared to open pollination, whereas the exclusion of nocturnal pollinators did not. There was significant selection on traits expected to influence pollinator attraction and pollination efficiency in both the diurnal and nocturnal pollination treatment. The relative strength of selection among plants exposed to diurnal and nocturnal visitors varied among traits and populations, but the direction of selection was consistent. The results suggest that diurnal pollinators are more important than nocturnal pollinators for seed production in the study populations, but that both categories contribute to selection on floral morphology. The study illustrates how experimental manipulations can link specific categories of pollinators to observed selection on floral traits, and thus improve our understanding of how species interactions shape patterns of selection.     

Parasitic flies (Diptera: Phoridae) influence foraging rhythms and caste division of labor in the leaf-cutter ant,Atta cephalotes (Hymenoptera: Formicidae)

Summary Three lines of evidence, including interspecific comparisons, temporal division of foraging between size castes, and experimental manipulations, suggest that the diurnal parasitoid Neodohrniphora curvinervis (Diptera: Phoridae) influences both the caste sizes and numbers of leaf-cutter ants (Atta cephalotes) that leave their underground nests to collect leaves. At Parque Nacional Corcovado in Costa Rica, A. cephalotes was attacked by Neodohrniphora during the daytime, and foraged less during the day than at night; a closely related ant at the same site, A. colombica, had no phorid parasites and foraged exclusively during the day. Most daytime foragers of A. cephalotes were smaller than the lower size threshold for attack by Neodohrniphora, while nocturnal foragers, active when parasitoids were absent, were both larger than this threshold and within the energetically optimal size range for foraging. When I supplied artificial lighting to allow phorids to hunt at A. cephalotes colonies past dusk, ants foraged less than when light was provided but flies were removed. The influence of Neodohrniphora on the foraging activity of A. cephalotes may explain why investigations focusing on abiotic factors have largely failed to discover what drives this ant's daily foraging cycles, and suggests that forager sizes are influenced not only by energetic efficiency, but also by the threat of parasitism.     

Linking fruit traits to variation in predispersal vertebrate seed predation, insect seed predation, and pathogen attack

The importance of vertebrates, invertebrates, and pathogens for plant communities has long been recognized, but their absolute and relative importance in early recruitment of multiple coexisting tropical plant species has not been quantified. Further, little is known about the relationship of fruit traits to seed mortality due to natural enemies in tropical plants. To investigate the influences of vertebrates, invertebrates, and pathogens on reproduction of seven canopy plant species varying in fruit traits, we quantified reductions in fruit development and seed germination due to vertebrates, invertebrates, and fungal pathogens through experimental removal of these enemies using canopy exclosures, insecticide, and fungicide, respectively. We also measured morphological fruit traits hypothesized to mediate interactions of plants with natural enemies of seeds. Vertebrates, invertebrates, and fungi differentially affected predispersal seed mortality depending on the plant species. Fruit morphology explained some variation among species; species with larger fruit and less physical protection surrounding seeds exhibited greater negative effects of fungi on fruit development and germination and experienced reduced seed survival integrated over fruit development and germination in response to vertebrates. Within species, variation in seed size also contributed to variation in natural enemy effects on seed viability. Further, seedling growth was higher for seeds that developed in vertebrate exclosures for Anacardium excelsum and under the fungicide treatment for Castilla elastica, suggesting that predispersal effects of natural enemies may carry through to the seedling stage. This is the first experimental test of the relative effects of vertebrates, invertebrates, and pathogens on seed survival in the canopy. This study motivates further investigation to determine the generality of our results for plant communities. If there is strong variation in natural enemy attack among species related to differences in fruit morphology, then quantification of fruit traits will aid in predicting the outcomes of interactions between plants and their natural enemies. This is particularly important in tropical forests, where high species diversity makes it logistically impossible to study every plant life history stage of every species.     

Dispersal of seeds by the tropical sea breeze

Given the dependence of most wind-pollinated and wind-dispersed species on low relative humidity (RH) for abscission, and the minimization of RH in the early afternoon, there ought to be a marked directional bias in seed dispersal at sites with a strong local diurnal circulation. We filmed the abscission of seeds of five wind-dispersed tropical species near the coast of Mexico (Jalisco). We found that (1) most abscission occurred during the period from 10:00 to 17:00 hours; (2) there was a strong bias for landward dispersal due to the midday sea breeze; (3) the little nocturnal dispersal that occurred was toward the sea (due to the night land breeze); (4) there was no abscission in the absence of wind (i.e., indoors) except for one species; and (5) holding relative humidity constant, the proportion of seeds diurnally abscising is strongly correlated with horizontal wind speed. We predict that a similar bias (this time for upslope dispersal) for xerochastic dispersal will occur in complex terrain. We conclude that in coastal and mountainous terrain, dispersal models (and inverse modeling efforts) must include a directionality term.     

The good and the bad stopover: behaviours of migrant reed warblers at two contrasting sites

Successful migration for passerine birds depends largely on the quality of stopover habitats, but we still lack complete knowledge of how migrants search for habitats en route and how they behave when landing at poor quality stopover sites. We compared the distance of exploratory movements and stopover durations of the reed warbler Acrocephalus scirpaceus, a reedbed habitat specialist, released at suitable (reed bed) and unsuitable (sand dune) stopover sites. Birds tape-lured during nocturnal migration to a sand dune were captured, radio-tagged, released and tracked at two sites of contrasting habitat quality. Lean birds were found to move further in the dunes (max. 300 m) than in reeds (max. 200 m), whereas ‘fat’ individuals at both sites remained stationary. Birds spent just 1 day in the dunes and up to 13 days in the reeds. Our results suggest that some nocturnal migrants with restricted diurnal exploratory movements depend on stopover site selection when ceasing nocturnal flight.     

Ants as a Measure of Effectiveness of Habitat Conservation Planning in Southern California

Abstract: In the United States multispecies habitat conservation plans were meant to be the solution to conflicts between economic development and protection of biological diversity. Although now widely applied, questions exist concerning the scientific credibility of the conservation planning process and effectiveness of the plans. We used ants to assess performance of one of the first regional conservation plans developed in the United States, the Orange County Central‐Coastal Natural Community Conservation Plan (NCCP), in meeting its broader conservation objectives of biodiversity and ecosystem‐level protection. We collected pitfall data on ants for over 3 years on 172 sites established across a network of conservation lands in coastal southern California. Although recovered native ant diversity for the study area was high, site‐occupancy models indicated the invasive and ecologically disruptive Argentine ant (Linepithema humile) was present at 29% of sites, and sites located within 200 m of urban and agricultural areas were more likely to have been invaded. Within invaded sites, native ants were largely displaced, and their median species richness declined by more than 60% compared with uninvaded sites. At the time of planning, 24% of the 15,133‐ha reserve system established by Orange County NCCP fell within 200 m of an urban or agricultural edge. With complete build out of lands surrounding the reserve, the proportion of the reserve system vulnerable to invasion will grow to 44%. Our data indicate that simply protecting designated areas from development is not enough. If habitat conservation plans are to fulfill their conservation promise of ecosystem‐level protection, a more‐integrated and systematic approach to the process of habitat conservation planning is needed.     

Relative intensity of predation on the French grunt, Haemulon flavolineatum, during diurnal, dusk, and nocturnal periods on a coral reef

Established hypotheses state that the rate of predation on coral reef fish should be highest during crepuscular periods (dawn and dusk) intermediate diurnally, and lowest nocturnally. We examined the relative risk of predation on juvenile French grunts (Haemulon flavolineatum Desmarest) during diurnal, dusk, and nocturnal periods on the fore- and back-reef at Teague Bay, U.S. Virgin Islands in July and August 1996. Tethering-devices recorded the exact amount of time between attaching a prey fish to its tether and subsequent predation on the prey fish. As tethering of prey usually inflates the actual rate of predation, times from our tethering devices were used to establish only the relative predation risk among treatments. During 3-h diurnal and nocturnal tethering experiments, relative predation was significantly higher during the nocturnal period, and differences between side of reef were not significant. In 30-min tethering experiments, which included all three time periods, the relative predation risk was significantly higher during dusk and nocturnal periods than during the diurnal period. Relative predation was not significantly different between the dusk and nocturnal periods, or between side of reef during any time period. The unexpected finding that the diurnal period had the lowest relative risk of predation indicates that the timing of predation events on reefs, as well as the adaptive reasons for nocturnal larval settlement, may need to be re-examined. Received: 11 February 1997 / Accepted: 21 October 1998     

Spatial and temporal strategies in the surface activity of some sandy beach arthropods living along the French Atlantic Coast

During a field study on the eco-ethology of sandy beach arthropods conducted along the French Atlantic Coast it emerged thatTylos europaeus, Phaleria cadaverina andLabidura riparia were among the most abundant species that lived in the eulittoral. Tetradirectional pitfall traps, placed from the base of the dune to the tidal limits, were used. The traps intercepted surface-active arthropods, and the number of traps varied according to the levels of the tides. Data were recorded hourly during two periods corresponding to a spring tide and the following neap tide. At the saine time the most important environmental parameters were registered each hour. The results indicate that all three species show prevalently nocturnal surface activities. ForT. europaeus andPh. cadaverina an influence of the synodic and tidal phases was found on the zonation of the surface activity and on the extent of movement. The latter was due to the greater or smaller distances that the isopods and tenebrionids had to move from the diurnal burrowing sites to the foraging zones. ForL. riparia no variations in the total mean zonations were found between the two synodic phases, and only at neap tide were seaward, followed by landward movements found. Multiple regression analysis showed that the environmental parameters registered in this period were never limiting, as threshold values in this case probably were not reached.     

Distribution,habitat preferences and behaviour of the critically endangered Maltese top-shell <Emphasis Type="Italic">Gibbula nivosa</Emphasis> (Gastropoda: Trochidae)

The Maltese top-shell, Gibbula nivosa Adams 1851, is a critically endangered marine trochid gastropod endemic to the Maltese Islands (central Mediterranean). Previously unrecorded live since 1981, rediscovery of this species in 2006 has enabled the collection of basic biological data for the first time, based on a population in Sliema Creek. The field distribution and habitat preferences of G. nivosa were investigated via stratified random sampling, while aspects of its behaviour were studied through laboratory experiments. Gibbula nivosa occurred primarily in near-homogeneous or homogeneous infralittoral cobble/pebble beds, at an overall density of 17.6 ± 25.0 SD individuals m⁻²; however, large spatial and temporal variations in abundance were recorded. The snails did not exhibit gregarious behaviour and the slightly clumped dispersion pattern noted in the field appeared to result from patchiness of the habitat. Recruitment was observed in post-summer months, which, together with data on shell-size distributions, suggests that G. nivosa spawns in early summer and attains adult size (maximum shell width ~9 mm) in less than 1 year. Gibbula nivosa showed a circadian activity pattern with nocturnal foraging involving indiscriminate ingestion of particles browsed off the substratum; observations on responses to four potential predators suggest that nocturnal activity may have evolved in response to diurnal predation. Historical records of snail occurrence and the present study suggest that population declines of G. nivosa in the past were probably related to habitat alteration as a result of anthropogenic pressure.     

Geographic variations in Helleborus foetidus elaiosome lipid composition: implications for dispersal by ants

Summary. Geographic variations in the correspondence between diaspore phenotypes and disperser behavior are thought to determine the evolution of plant-animal dispersal mutualisms. Helleborus foetidus is a widely distributed plant in Western Europe, which seeds bear a lipid rich elaiosome attracting ant dispersers. Laboratory cross-tests were conducted to check the correspondence between diaspore phenotypes and ant preference in two localities of the Iberian Peninsula, Caurel and Cazorla, separated by 750 km. Diaspores from Caurel were systematically preferred to those from Cazorla by Formica lugubris (the major disperser at Caurel), and Aphaenogatser iberica and Camponotus cruentatus (both major dispersers at Cazorla). Further bioassays conducted on A. iberica only showed that differences in elaiosome traits were sufficient to explain ant preference. Separation of the lipid fractions composing the elaiosome revealed that triglycerides, diglycerides and free fatty acids were all dominated by oleic acid. The elaiosomes from Caurel contained relatively more free oleic acid but were less concentrated in linoleyl-containing triglycerides, free palmitic acid and free linoleic acid than those from Cazorla. The three lipid fractions were attractive to ants but dummies soaked with the free fatty acids extracted from Caurel were preferred to those from Cazorla. Taken together, these results reinforce the idea that oleic acid is a major releaser of seed collection by ants and suggest that geographic variations in free fatty acid composition affect the probability of diaspore removal by ants which in turn potentially determine plant demography.     

Preparation and characterisation of green clay-polymer nanocomposite for heavy metals removal

ABSTRACT

Natural polymer Moringa oleifera seed as coagulant and bentonite clay as adsorbent were used for preparing novel composite coagulant. Results obtained from FTIR, SEM, TEM and P-XRD show that the bentonite clay and M. oleifera seed biopolymers physico-chemically interact with each other during the preparation of clay-polymer composite. The FTIR results show that the major functional groups present in bentonite clay and M. oleifera seed are integrated at nano levels in the novel composite to remove heavy metals from aqueous systems. The coagulo-adsorption using clay-polymer composite may be used for the adsorption of heavy metals ions from the aqueous systems. It becomes possible due to the structural characteristics of the clay crystallites together with the functional attraction of the biopolymer and it results in the formation of clay-polymer metal complexes. The clay-polymer nano-composite of M. oleifera seed and bentonite clay showed considerable cadmium, chromium and lead removal property.     

Incorporating animal behavior into seed dispersal models: implications for seed shadows

Seed dispersal fundamentally influences plant population and community dynamics but is difficult to quantify directly. Consequently, models are frequently used to describe the seed shadow (the seed deposition pattern of a plant population). For vertebrate-dispersed plants, animal behavior is known to influence seed shadows but is poorly integrated in seed dispersal models. Here, we illustrate a modeling approach that incorporates animal behavior and develop a stochastic, spatially explicit simulation model that predicts the seed shadow for a primate-dispersed tree species (Virola calophylla, Myristicaceae) at the forest stand scale. The model was parameterized from field-collected data on fruit production and seed dispersal, behaviors and movement patterns of the key disperser, the spider monkey (Ateles paniscus), densities of dispersed and non-dispersed seeds, and direct estimates of seed dispersal distances. Our model demonstrated that the spatial scale of dispersal for this V. calophylla population was large, as spider monkeys routinely dispersed seeds >100 m, a commonly used threshold for long-distance dispersal. The simulated seed shadow was heterogeneous, with high spatial variance in seed density resulting largely from behaviors and movement patterns of spider monkeys that aggregated seeds (dispersal at their sleeping sites) and that scattered seeds (dispersal during diurnal foraging and resting). The single-distribution dispersal kernels frequently used to model dispersal substantially underestimated this variance and poorly fit the simulated seed-dispersal curve, primarily because of its multimodality, and a mixture distribution always fit the simulated dispersal curve better. Both seed shadow heterogeneity and dispersal curve multimodality arose directly from these different dispersal processes generated by spider monkeys. Compared to models that did not account for disperser behavior, our modeling approach improved prediction of the seed shadow of this V. calophylla population. An important function of seed dispersal models is to use the seed shadows they predict to estimate components of plant demography, particularly seedling population dynamics and distributions. Our model demonstrated that improved seed shadow prediction for animal-dispersed plants can be accomplished by incorporating spatially explicit information on disperser behavior and movements, using scales large enough to capture routine long-distance dispersal, and using dispersal kernels, such as mixture distributions, that account for spatially aggregated dispersal.     

Diurnal vertical migration of an estuarine cladoceran,Podon polyphemoides,in the Chesapeake Bay

Populations of the marine cladoceran Podon polyphemoides (Leuckart) in the Chesapeake Bay (USA) zooplankton typically occur in large patches, extending the length of the estuary. These patches are characterized by well defined maxima, which do not move seaward during a given season. During the day, the bulk of the population is situated in the upper part of the two-layered estuarine circulation, where it is subjected to transport by net non-tidal seaward current. One mechanism for maintaining the population within the estuary is provided by the diurnal behavior of the podonids, which migrate to deeper water during the night. The landward advection of the deeper currents, thus, helps to counterbalance seaward population loss during the day. The extent of the vertical movement of the population can be modified by various environmental factors, but a suggestive correlation with light was found. It is thought that the downward component of the migration is due to passive sinking.     

Room to move? Threatened shorebird habitat in the path of sea level rise—dynamic beaches, multiple users, and mixed ownership: a case study from Rhode Island, USA

Accelerated sea level rise (slr) is expected to transform vulnerable Atlantic coastal habitats in the United States during this century. Low-elevation sandy beaches, important nesting habitat for the continued recovery of the federally threatened piping plover (Charadrius melodus) in Rhode Island, are especially vulnerable. These beaches, under a mix of private and public ownership, exist in a heterogeneous landscape of dunes, rocky headlands, salt ponds, and heavily developed areas. Understanding the extent to which piping plover nesting beaches can retreat landward under projected slr is important for prioritizing future conservation actions across multiple jurisdictions. This research examines habitat change in response to slr (0.5, 1.0, and 1.5 m) under stationary and migration beach responses and whether development blocks habitat migration for five productive piping plover nesting beaches. We found that under the stationary habitat model, all beaches lose area under all slr scenarios. For the habitat migration model, future habitat availability differs by beach depending on elevation, landward topography, and presence of development. However, across the majority of beach area, piping plover habitat will be able to migrate landward if unconstrained by future development. A coalition of public and private stakeholders already supports management efforts to help ensure plover population recovery in the area. With emerging habitat change prediction tools, these stakeholders and other partners can engage in innovative, local-level planning needed to protect wildlife habitat and commercial, residential, and infrastructure assets from sea level rise.     

Extremely high mating frequency in the Florida harvester ant (<Emphasis Type="Italic">Pogonomyrmex badius</Emphasis>)

High effective multiple queen mating is a rare but taxonomically widespread phenomenon in eusocial Hymenoptera that has arisen convergently in several taxa. In ants, high effective mating frequencies have been confirmed in only two clades: the higher leaf-cutters (Atta, Acromyrmex) and the Pogonomyrmex seed harvesters. We analysed polyandry in Pogonomyrmex badius, which has a life-history unique within the genus, and report the highest numerical mating frequencies thus far recorded in ants. We also show that P. badius is characterized by one of the highest effective mating frequencies hitherto found in ants. It is now clear that all major sub-clades of Pogonomyrmex sensu stricto exhibit high levels of polyandry. Therefore, multiple mating must have arisen early in the evolution of the genus, and may have constituted a mechanism to increase offspring variability for queens that were confronted with increasingly complex levels of organization. Too few congeners have been investigated by the same method to be certain that polyandry in P. badius is really higher than in the rest of the genus. If so, research should concentrate on a possible link between high queen mating frequency and the distinct caste system found in P. badius.Communicated by L. Keller     

Burrowing behaviour inferred from feeding traces of the opheliid polychaete <Emphasis Type="Italic">Euzonus</Emphasis> sp. as response to beach morphodynamics

The distribution of the opheliid polychate Euzonus sp. and the orientation of its feeding traces were examined in a wave-dominated Pacific sandy beach of central Japan by periodical sampling from June to December 2006. This species occurred constantly within sediments at mid-intertidal level, although its horizontal distributions were variable depending on changes of the beach profiles. These facts indicate that Euzonus sp. moved horizontally seaward or landward within the substratum in response to the shifting beach face due to changes of wave conditions from the ocean. By analysing the orientation of feeding traces, the adaptive burrowing behaviour of this species in response to beach topographical changes was revealed: Euzonus worms burrowed horizontally to various directions in a relatively stable condition of beach topography, whereas they moved preferentially landward under a heavy erosion of the beach face by a storm. These findings indicate that the burrowing behaviour prevents Euzonus sp. from excessive burial and washing out due to the beach morphodynamics.     

Feeding over the 24-h cycle: dietary flexibility of cathemeral collared lemurs (<Emphasis Type="Italic">Eulemur collaris</Emphasis>)

Animals show specific morphological, physiological and behavioural adaptations to diurnal or nocturnal activity. Cathemeral species, i.e. animals with activities distributed over the 24-h period, have to compromise between these specific adaptations. The driving evolutionary forces and the proximate costs and benefits of cathemerality are still poorly understood. Our goal was to evaluate the role of predator avoidance, food availability and diet quality in shaping cathemeral activity of arboreal mammals using a lemur species as an example. For this, two groups of collared lemurs, Eulemur collaris, were studied for 14 months in the littoral forest of southeastern Madagascar. Data on feeding behaviour were collected during all-day and all-night follows by direct observation. A phenological transect containing 78 plant species was established and monitored every 2 weeks to evaluate food availability during the study period. Characteristics of food items and animal nutritional intake were determined via biochemical analyses. The ratio of diurnal to nocturnal feeding was used as response variable in the analyses. The effects of abiotic environmental variables were removed statistically before the analyses of the biotic variables. We found that diurnal feeding lasted longer during the hot–wet season (December–February), whereas nocturnal feeding peaked during the hot–dry and cool–wet seasons (March–August). Although the lemurs foraged mostly in lower forest strata during daylight and used emergent trees preferably at night, the variables which measured animal exposure to birds of prey failed to predict the variation of the ratio of diurnal/nocturnal feeding. Ripe fruit availability and fiber intake are the two variables which best predicted the annual variation of the lemur diurnality. The data indicate that feeding over the whole 24-h cycle is advantageous during lean periods when animals have a fibre-rich, low-quality diet.