Foraging behaviour of harvesting ants determines seed removal and dispersal

Harvesting ants can affect the regeneration of plants through at least two different processes: seed removal and seed dispersal. We analyse the role of different foraging strategies of ants on patterns of seed removal and dispersal by three Messor species with considerable differences in their foraging systems. Messor capitatus workers rarely leave the nest in well-formed columns, while the other two species form foraging trails, with M. bouvieri forming temporary trails and M. barbarus foraging on a stable system of permanent foraging trails. Overall seed intake of M. capitatus colonies is considerably less than that of the two group-foraging species. There are also differences in the size of seeds collected: M. barbarus and M. capitatus harvest similar amounts of large and small seeds, while M. bouvieri harvests small seeds more intensely than large ones, due to the smaller size of the worker caste. The three Messor species differ in the percent of seed dropping of the different seed type and in the seed dispersal distance. Moreover, M. bouvieri and M. capitatus redistributed dropped seeds preferentially in bare soil and low sparse vegetation habitats, while M. barbarus redistributed seeds mainly in the high vegetation habitat. These results show that the foraging systems of these harvesting ants determine different patterns of seed removal and dispersal and, thus, affect the abundance and redistribution of seeds in the area.     

Connecting fruit production to seedling establishment in two co-occurring <Emphasis Type="Italic">Miconia</Emphasis> species: consequences of seed dispersal by birds in upper Amazonia

This study investigated links between seed production by two species of Miconia (Melastomataceae), whose seeds are dispersed by birds, and later stages of recruitment in lowland forests of eastern Ecuador. Seed dispersal and survival in later stages are crucial for understanding and predicting patterns of plant population dynamics as well as for understanding patterns of diversity in tropical forests. A major goal was to determine if the spatial template of seed deposition established by birds predicted probability of recruitment. We used observational and experimental approaches to compare patterns of recruitment in Miconia fosteri and M. serrulata. We calculated probabilities of transition between successive stages of recruitment for each species in three habitats. The number of plants with fruit, number of fruits removed, and, to a lesser extent, patterns of seed deposition varied between species and among habitats, whereas seed survival, germination, and establishment showed little variation among habitats. The location of seed deposition directly influenced the cumulative probabilities of survival. Among-habitat differences in the probabilities of recruitment set by seed deposition were not modified by later stages, although probability of recruitment was 2.5 times higher for M. serrulata than for M. fosteri after 1 year. The more critical stages for recruitment were seed removal and deposition. Our results from multiple life-cycle stages suggest that habitat associations among plants that reach reproductive maturity become established at early life stages and were mostly a consequence of seed dispersal by birds. These results differ from those obtained in temperate zones and suggest fundamental differences in the importance of recruitment processes. Dispersers, such as manakins, play significant roles in recruitment and population dynamics of M. fosteri, M. serrulata and numerous other understory plants of Neotropical forests. Their role in plant recruitment could be much greater than previously considered in megadiverse tropical forests. Thus, loss of dispersers could have long-term and far-reaching implications for maintenance of diversity.     

Influence of temporal fluctuations in seed abundance on the foraging behaviour of harvester ants (Pogonomyrmex spp.) in the central Monte desert,Argentina

The simultaneous study of the temporal dynamics of foraging behaviour, diet and seed abundance is essential to assess the way in which resources affect the behaviour and ecology of harvester ants. Here, we evaluate how fluctuations in grass seed abundance during three consecutive growing seasons influenced the foraging behaviour and diet of the harvester ants Pogonomyrmex rastratus, P. mendozanus and P. inermis in the central Monte desert, Argentina. Seed abundance of the most consumed grasses varied greatly through ant activity season, and ants altered their foraging behaviour in response to those changes. Foragers spent more time travelling and searching for food, and their foraging trips took longer during the low seed availability season. Foraging distance was very similar among species and, contrary to our expectations, did not vary between seasons. Foraging success of P. rastratus and P. inermis increased during the high availability season. This matched the seasonal pattern of foraging activity, suggesting that colonies may detect seed abundance and regulate their foraging effort with the rate of forager success. Although grass seeds were the main item in the diet of the three species, P. mendozanus, and to a lesser extent P. rastratus, turned more generalist when grass seeds were scarce. In contrast, P. inermis showed a very narrow diet breadth, only harvesting grass seeds in both seasons. Our results indicate the relevance of seed availability on foraging behaviour of harvester ants, which should be taken into account when predicting and evaluating the effect of ants on seed resources as well as numerical responses of harvester ant populations to the temporal and spatial variations in grass seed abundance.     

Spatial patterns of seed predation by harvester ants (<Emphasis Type="Italic">Messor</Emphasis> Forel) in Mediterranean grassland and scrubland

Summary. Seed predation by harvester ants is one of the main processes involved in the seed bank dynamics of Mediterranean grassland and scrubland. We investigated the spatial patterns of seed predation by Messor spp. in these systems from a multi-level approach (nest-site selection, foraging patch selection and seed predation risk), using ten experimental plots (20 m × 40 m) set in central Spain. All habitats in grasslands are dominated by M. barbarus, while in scrubland this species shares seed exploitation with M. bouvieri. In this type of system, the nest-hole and foraging territory distribution of the two species show a certain degree of segregation. Messor bouvieri focuses on the exploitation of dry scrub, while M. barbarus also forages on subhumid and humid patches. We interpret this as a consequence of the different foraging behaviour and nest site requirements of the two species. In general, the spatial distribution of seed predation risk reflects the patterns detected for the nest-holes and foraging areas, although scrubland has a more heterogeneous predation pattern than grassland.     

Avian seed dispersal and seedling distribution of the endangered tree species,Taxus chinensis,in patchy habitats

Background: There is limited understanding about bird dispersal behaviour and seedling distribution of endangered tree species in patchy environments, although these processes are important for plant species persistence.

Aims: We tested how patch features affected bird behaviour and seed dispersal, and thus seedling distribution of the endangered Chinese yew tree (Taxus chinensis).

Methods: In the present study, we combined field data of bird dispersal behaviour and GIS-based information to elucidate the influence of spatial features of habitat patches on bird dispersal behaviour, and the resulting effects on the seedling distribution of the endangered Chinese yew in two patchy habitats.

Results: Our results showed that the only seed source patch could attract eight bird species for dispersal at the two sites. Post-foraging movements of bird dispersers was strongly related to both topography and the relative locations of habitat patches. Yew seedlings aggregated only at the seed source and bamboo recruitment patches, which was affect by both the spatial distribution of recruitment patches and patch use by dispersers.

Conclusions: Our results emphasise that bamboo patches in both patchy environments provide the necessary conditions for germination of yew seeds, and the post-foraging behaviour of dispersers determines seed deposited in these patches. Our study highlights the importance of the dispersal behaviour of frugivorous birds in the successful regeneration and colonisation of yew populations in patchy habitats.     

Relationships between intra‐specific variation in seed size and recruitment in four species in two contrasting habitats

Large seeds contain more stored resources, and seedlings germinating from large seeds generally cope better with environmental stresses such as shading, competition and thick litter layers, than seedlings germinating from small seeds. A pattern with small‐seeded species being associated with open habitats and large‐seeded species being associated with closed (shaded) habitats has been suggested and supported by comparative studies. However, few studies have assessed the intra‐specific relationship between seed size and recruitment, comparing plant communities differing in canopy cover. Here, seeds from four plant species commonly occurring in ecotones between open and closed habitats (Convallaria majalis, Frangula alnus, Prunus padus and Prunus spinosa) were weighed and sown individually (3200 seeds per species) in open and closed‐canopy sites, and seedling emergence and survival recorded over 3 years. Our results show a generally positive, albeit weak, relationship between seed size and recruitment. In only one of the species, C. majalis, was there an association between closed canopy habitat and a positive seed size effect on recruitment. We conclude that there is a weak selection gradient favouring larger seeds, but that this selection gradient is not clearly related to habitat.     

Ant-mediated expansion of an obligate seeder species during the first years after fire

Most obligate seeder species build up a soil seed bank that is associated with massive seed germination in the year immediately after a fire. These species are also shade‐intolerant and disappear when vegetation cover closes, creating unsuitable conditions for seedling recruitment. The only way for these plants to expand their populations is when habitats suitable for seedling recruitment arise (i.e. in years immediately after a fire). However, short primary seed dispersal of obligate seeders does not allow these plants to colonise the suitable habitats, and these habitats can only be colonised by secondary seed dispersion. We hypothesised that Fumana ericoides, an obligate‐seeding small shrub, not only establishes abundantly in the first year after fire, but also expands its local range in the following years due to secondary dispersal by ants while suitable habitats are still available. We tested this hypothesis using experimental studies and a simulation model of potential population expansion in a recently burned area. Results showed that F. ericoides not only established prolifically in the year immediately after fire, but was also able to recruit new individuals and expand its population in the years following the fire, despite a low germination rate and short primary seed dispersal. Ant‐mediated seed dispersal and availability of suitable habitats were key factors in this phenomenon: ants redistributed seeds in suitable habitats while they were available, which accelerated the expansion of F. ericoides because new plants established far away from the core population.     

How an invasive alga species (<Emphasis Type="Italic">Caulerpa taxifolia</Emphasis>) induces changes in foraging strategies of the benthivorous fish <Emphasis Type="Italic">Mullus surmuletus</Emphasis> in coastal Mediterranean ecosystems

The foraging behaviour variability of three striped red mullet (Mullus surmuletus L. 1758) populations with respect to the vegetation cover was examined along the French Mediterranean coasts. We tested both the carrying capacity of different habitats and the hypothesis of a food segregation by the invasive alga Caulerpa taxifolia (Vahl) C. Agardh, on the functioning of benthic fish populations. The results indicated a significant site effect on M. surmuletus foraging behaviour and movements. The vegetation cover may play an important role in modifying the striped red mullet cost of foraging (sampling effort) and thus its strategy of prey capture. As long as the cover of marine phanerogams and/or macrophyte algae (Caulerpaceae) increases, the foraging budget and the distances covered in search of prey decrease significantly. Similarly, the striped red mullet increases the prospected sample periods to augment its foraging success, swimming above the bottom. These changes are related to the development of a dense superficial network of plagiotropic rhizomes and stolons, and to the reduction of space between fronds that limits the accessibility to resources and may increase intra-specific food competition. The role of C. taxifolia does not differ from that of other marine phanerogams but induces significant changes in the structure of Mullidae populations at the local level. The rapid expansion of Caulerpaceae in the Mediterranean Sea could constitute a real threat for the balance of the marine coastal biodiversity and the ecology of M. surmuletus, which is considered a flagship species for coastal Mediterranean demersal fisheries.     

Pollination and seed dispersal of Melocactus ernestii Vaupel subsp. ernestii (Cactaceae) by lizards: an example of double mutualism

Recent studies show that the mutualistic role of lizards as pollinators and seed dispersers has been underestimated, with several ecological factors promoting such plant–animal interactions, especially on oceanic islands. Our aim is to provide a quantitative assessment of pollination and seed dispersal mutualisms with lizards in continental xeric habitats. We carried out focal observations of natural populations of Melocactus ernestii (Cactaceae) in the Caatinga, a Brazilian semiarid ecosystem, in order to record the frequency of visits, kind of resource searched and behaviour of visiting animals towards flowers and/or fruits. We made a new record of the lizard Tropidurus semitaeniatus foraging on flowers and fruits of M. ernestii. During the search for nectar, T. semitaeniatus contacted the reproductive structures of the flowers and transported pollen attached to its snout. Nectar production started at 14:00 h, with an average volume of 24.4 μl and an average concentration of solutes of 33%. Approximately 80% of the seeds of M. ernestii found in the faeces of T. semitaeniatus germinated under natural conditions. The roles of T. semitaeniatus as pollinator and seed disperser for M. ernestii show a clear relationship of double mutualism between two endemic species, which may result from the environmental conditions to which both species are subject. Seasonality, low water availability and arthropod supply in the environment, high local lizard densities, continuous nectar production by the flower and fruits with juicy pulp may be influencing the visits and, consequently, pollination and seed dispersal by lizards in this cactus.     

斑块生境中啮齿类和蚁类动物对南方红豆杉地面种子的搬运

南方红豆杉(Taxus chinensis)在我国南方地区多分布在村落附近的斑块生境中,成熟后的种子直接落在母树下或被鸟类等动物搬运至其他斑块中。林下地面种子会被地面活动的动物搬运,继而影响种子命运及种群更新。于2016年、2017年南方红豆杉果期,在浙江天目山一个红豆杉种群分布地(临安市桐坑村),采用野外种子摆放实验的方法研究了动物对地面种子的搬运情况。结果表明:啮齿类动物取食是南方红豆杉地面种子消失的主要原因,但种子消失率在斑块间及边缘生境中的差异较大。母树林和竹林是啮齿类动物的主要觅食生境,两种生境的动物取食率明显高于山核桃种植园。边缘生境成为啮齿类斑块间移动的通道,而非觅食场所。4种啮齿类动物中,淡腹松鼠(Callosciurus pygerythrus)在母树林斑块以外的生境中出现频率最高,而其他地面搬运者回避利用山核桃种植园,仅在母树林和竹林斑块中搬运南方红豆杉地面种子。可见,斑块生境中动物改变了南方红豆杉地面种子的空间沉积格局,进而影响植物种群的更新。     

Foraging habitats of bats in southern Finland

We determined the foraging habitats of the northern batEptesicus nilssonii (Keyserling et Blasius, 1839), Brandt’s batMyotis brandtii (Eversmann, 1845), whiskered batMyotis mystacinus (Kuhl, 1819), Daubenton’s batMyotis daubentonii (Kuhl, 1819) and brown long-eared batPlecotus auritus (Linnaeus, 1758) in southern Finland. Among these species, we compared the diversities of foraging habitats, linear feature preference and the bats’ tendencies to forage simultaneously.Eptesicus nilssonii was the most opportunistic, foraging in a wide range of habitats.Myotis daubentonii (94%) foraged mainly on water habitats, whileM. brandtii/mystacinus (89%) andP. auritus (66%) foraged mainly in forest habitats. The diversities of foraging habitats used byE. nilssonii andP. auritus were higher than those ofM. brandtii/mystacinus andM. daubentonii. Both E.nilssonii andP. auritus foraged mostly alone or in small numbers, whileM. brandtii/mystacinus tended to gather in large numbers to forage in the same habitat. Half ofE. nilssonii and 46% ofM. daubentonii foraged over linear features, while other species did not use linear features to such an extent. Management and conservation of foraging habitats are needed especially forM. brandtii/mystacinus andM. daubentonii, which are more specialized thanE. nilssonii and P. auritus.     

Variation in resource size distribution around colonies changes ant–parasitoid interactions

The distribution of resources within habitats affects species abundance, richness and composition, but the role of resource distribution in species interactions is rarely studied. In ant communities, changes in resource distribution within habitats may influence behavioral interactions because many ant species are specialized to efficiently harvest a subset of available resources. This study investigates whether interactions between the behaviorally dominant host ant Pheidole diversipilosa and its specialist parasitoid (Phoridae: Apocephalus orthocladus) depend on resource size distribution around the colony. Using in situ foraging arenas to manipulate parasitoid abundance and resource size distribution around colonies, we tested whether variation in resource size distribution allows P. diversipilosa to alter its foraging behavior in ways that lessen the impact of parasitoid attack. P. diversipilosa colonies do not lower the impact of parasitoid attack by increasing the number of workers foraging individually on small and widely dispersed resources. However, the presence of multiple large resources allows colonies to temporarily redistribute soldier ants from resources patrolled by parasitoids to other resources not patrolled by parasitoids, and to maintain soldier abundance at levels found in the absence of parasitoids. These results highlight the importance of placing behavioral interactions within the context of variation in resource distribution.     

Seed to seedling transitions in successional habitats across a tropical landscape

Recognition that tree recruitment depends on the balance between seed arrival and seedling survival has led to a surge of interest in seed‐dispersal limitation and seedling‐establishment limitation in primary forests. Virtually unaddressed are comparisons of this balance in mature and early successional habitats. We assessed seed rain and seedling recruitment dynamics of tree species in primary forest, secondary forest and pasture released from grazing in a tropical agricultural landscape. Seed to seedling ratios (seed effectiveness; Φ_i) for 43 species in southern Mexico determined differences in the extent to which seeds produced seedlings by habitat, life history, and dispersal mode. Reproductive potential as estimated by the transition from seed rain to seedling recruitment, differed by habitats, and varied dramatically by life history and dispersal mode. Expected recruit densities (E_it) were higher for animal‐dispersed than wind‐dispersed species, and for non‐pioneer than pioneer species. Non‐pioneers and animal‐dispersed species had higher expected relative recruit abundance (ε_it) in primary forest (median of 4 seeds recruit^?1) whereas in secondary forest wind‐dispersed pioneers had the highest expected relative recruit abundance (median of 16 seeds per recruit). In pastures, wind‐dispersed pioneer species were most successful with many more seeds per recruit (median of 291) than both forest habitats. Seeds per recruit (Φ_i) appeared to decrease with increase in seed mass for 43 species for which data were available (r = –0.55, P < 0.001). This was associated with a negative correlation of Φ_i with seed size in primary forest (r = –0.50, P = 0.08 for 13 species); Φ_i was not correlated with seed size in secondary forest (n = 16) or pasture (n = 14). Metrics of seeds per recruit, expected recruit density and expected relative recruit abundance dramatically illustrate differences in barriers to recruitment in successional habitats.     

Influence of temporal fluctuations in seed abundance on the diet of harvester ants (Pogonomyrmex spp.) in the central Monte desert,Argentina

Harvester ants usually go through temporal fluctuations in environmental seed abundance and composition which could influence their behaviour and ecology. The aim of this study was to evaluate how these fluctuations influence the diet of Pogonomyrmex rastratus, P. pronotalis and P. inermis (Hymenoptera, Formicidae) in the central Monte desert during three consecutive growing seasons. Although seeds were the main item in the diet, these ants turned more generalist when seed abundance of the most consumed species (grasses Aristida spp., Trichloris crinita, Pappophorum spp., Digitaria californica and Stipa ichu) was low. Accordingly, diversity of items in the diet decreased with seed abundance in a logarithmical fashion, showing higher foraging efficiency for seeds at higher seed abundance. Seed diversity, however, was not related to seed abundance as ants always included several species in their diet, with alternating prevalence. The proportion of the most consumed species increased logarithmically in the diet of P. rastratus and P. pronotalis along with their abundance in the environment probably as a consequence of diet switching (from forb and shrub seeds to grass seeds) and by an increase in foraging efficiency at higher seed densities. In contrast, foraging activity of P. inermis was very low at low seed abundance and its diet included only the five grasses. Among the most consumed species, proportion in the diet was not associated with relative abundance in the environment. Aristida spp., Pappophorum spp. and D. californica were overall highly selected. However, the flexibility in the diet of P. pronotalis and P. rastratus and the low foraging activity of P. inermis during periods of low resource abundance could attenuate potential top‐down effects in the central Monte desert. This study shows that bottom‐up effects are important in ant‐seed interactions and should be considered when predicting and evaluating ants' effects on seed resources.     

Floral display size in comfrey, Symphytum officinale L. (Boraginaceae): relationships with visitation by three bumblebee species and subsequent seed set

The fecundity of insect-pollinated plants may not be linearly related to the number of flowers produced, since floral display will influence pollinator foraging patterns. We may expect more visits to plants with more flowers, but do these large plants receive more or fewer visits per flower than small plants? Do all pollinator species respond in the same way? We would also expect foragers to move less between plants when the number of flowers per plant are large, which may reduce cross-pollination compared to plants with few flowers. We examine the relationships between numbers of inflorescence per plant, bumblebee foraging behaviour and seed set in comfrey, Symphytum officinale, a self-incompatible perennial herb. Bumblebee species differed in their response to the size of floral display. More individuals of Bombus pratorum and the nectar-robbing B.?terrestris were attracted to plants with larger floral displays, but B. pascuorum exhibited no increase in recruitment according to display size. Once attracted, all bee species visited more inflorescences per plant on plants with more inflorescences. Overall the visitation rate per inflorescence and seed set per flower was independent of the number of inflorescences per plant. Variation in seed set was not explained by the numbers of bumblebees attracted or by the number of inflorescences they visited for any bee species. However, the mean seed set per flower (1.18) was far below the maximum possible (4 per flower). We suggest that in this system seed set is not limited by pollination but by other factors, possibly nutritional resources.     

Electrocommunication behaviour during social interactions in two species of pulse‐type weakly electric fishes (Mormyridae)

This study compares electrocommunication behaviour in groups of freely swimming weakly electric fishes of two species, Marcusenius altisambesi and Mormyrus rume. Animals emitted variable temporal sequences of stereotyped electric organ discharges (EOD) that served as communication signals. While the waveform of individual signals remained constant, the inter‐discharge interval (IDI) patterns conveyed situation‐specific information. Both species showed different types of group behaviour, e.g. they engaged in collective (group) foraging. The results show that in each species, during different behavioural conditions (resting, foraging and agonistic encounters), certain situation‐specific IDI patterns occurred. In both species, neighbouring fishes swimming closely together interacted electrically by going in and out of synchronization episodes, i.e. periods of temporally correlated EOD production. These often resulted in echo responses between neighbours. During group foraging, fishes often signalled in a repetitive fixed order (fixed‐order signalling). During foraging, EOD emission rates of M. altisambesi were higher and more regular than those of M. rume. The two species also differed in the quantity of group behaviours with M. altisambesi being more social than M. rume, which was reflected in the lack of specific agonistic IDI patterns, more fixed‐order signalling and more communal resting behaviour in M. altisambesi.     

Scratching beneath the surface: Bandicoot bioturbation contributes to ecosystem processes

Animals that forage for food or dig burrows by biopedturbation can alter the biotic and abiotic characteristics of their habitat. The digging activities of such ecosystem engineers, although small at a local scale, may be important for broader scale landscape processes by influencing soil and litter properties, trapping organic matter and seeds, and subsequently altering seedling recruitment. We examined environmental characteristics (soil moisture content, hydrophobicity and litter composition) of foraging pits created by the southern brown bandicoot (Isoodon obesulus; Peramelidae), a digging Australian marsupial, over a 6‐month period. Fresh diggings typically contained a higher moisture content and lower hydrophobicity than undisturbed soil. A month later, foraging pits contained greater amounts of fine litter and lower amounts of coarse litter than adjacent undug surfaces, indicating that foraging pits may provide a conducive microhabitat for litter decomposition, potentially reducing litter loads and enhancing nutrient decomposition. We tested whether diggings might affect seedling recruitment (seed removal by seed harvesters and seed germination rates) by artificially mimicking diggings. Although there were no differences in the removal of seeds, seedling recruitment for three native plant species (Acacia saligna, Kennedia prostrata and Eucalyptus gomphocephala) was higher in plots containing artificial diggings compared with undug sites. The digging actions of bandicoots influenced soil moisture and hydrophobicity, the size distribution of litter and seedling recruitment at a local scale. The majority of Australian digging mammals are threatened, with many suffering substantial population and range contraction. However, their persistence in landscapes plays an important role in maintaining the health and function of ecosystems.     

Behavioural responses of the lizard Pedioplanis l. lineoocellata to overgrazing

We studied the effects of overgrazing on the foraging behaviour of the lizard Pedioplanis l. lineoocellata (Spotted Sand Lizard), a sit-and-wait forager, in habitats of differing vegetation states to determine the effects of habitat degradation on this species. At high grazing intensity where vegetation cover and diversity is low, the lizard P. lineoocellata moves more frequently, spends more time moving and covers larger distances than in habitats where vegetation cover and diversity is high. These behavioural changes in movement patterns can be explained by less abundant prey in habitats with low vegetation cover and diversity. Although morphology, phylogeny and physiology of P. lineoocellata should constrain the change in foraging behaviour, the species has modified its foraging strategy from sit-and-wait to actively foraging. We assume that this behavioural flexibility of P. lineoocellata is a buffer mechanism enabling the species to use and survive in degraded (unfavourable) habitats.     

Foraging behavior and morphology: seed selection in the harvester ant genus, Pogonomyrmex

Optimally foraging animals can be behaviorally or morphologically adapted to reduce the energetic and time costs of foraging. We studied the foraging behavior and morphology of three seed harvester ant species, Pogonomyrmex barbatus, P. desertorum, and P. occidentalis, to determine the importance of behavioral strategies and morphological features associated with load carriage in reducing the costs of foraging. We found that none of five morphological features we measured had a significant impact on seed selection. Also, body size did not influence running speed, an important variable in time costs of foraging. Temperature had the largest effect on running speed in these species. Our results show that these species have foraging strategies which minimize the time costs of traveling with seeds. We also describe a pattern where the running speed in individual-foraging species is less affected by increasing seed size than in trunk-trail foragers, when temperature and body mass are held constant. These results support previous work which showed that time costs are most important in seed selection for Pogonomyrmex, and suggest that central place foraging theory may need to accommodate variation in foraging strategy to more accurately predict optimal seed size selection in harvester ants. Received: 16 June 1997 / Accepted: 15 December 1997     

Coexistence and Habitat Preference of Two Honeyeaters and a Sunbird on Lombok,Indonesia

On the island of Lombok, Indonesia, three nectarivorous birds partially coexist: the two closely related and very similar Lichmera honeyeaters and a sunbird (Cinnyris jugularis). We investigated how these species segregated ecologically in areas where they coexisted by evaluating foraging visits and aggressive interactions at rich and poor nectar resources in different habitats (forest and open areas) along an altitudinal gradient (800–1600 m asl). The two honeyeaters were partially segregated by altitude. In the zone of overlap, Lichmera limbata dominated the richest forest resources, while Lichmera lombokia dominated the richest resources in open land. The sunbird, C. jugularis, was only observed in open habitats and mostly at poor resource patches. In the three‐species community in open habitats a dominance hierarchy was apparent with L. lombokia as the superior species and C. jugularis as the inferior species. Studies on how segregated species interact in their transition zone can help us to understand interactions between otherwise ecologically segregated species.