High seed dispersal rates in faunally intact tropical rain forest: theoretical and conservation implications

Dispersal limitation may promote high tree-species diversity in rain forest by slowing local competitive exclusion, but evidence is scarce. By comparing the species lists of tree plots with those of nested seedling plots in rain forest at Gunung Palung, we found that the proportion of local seedlings arising from active dispersal events (via wind or animals, but not simply dropped from parent trees) was high: 68% of species and 46% of the individual seedlings. Local seedling species richness was not limited by the local richness of adults. Using these data, a spatially explicit simulation model indicated that dispersal limitation may not contribute substantially to the maintenance of tree diversity in this forest. We predict that the loss of animal seed dispersers would reduce local species richness of seedlings by 60%. While this reduction could possibly increase the influence of dispersal limitation, it would interfere with other mechanisms maintaining diversity.     

Leaf density and a trade-off between load-size selection and recruitment behavior in the ant Atta cephalotes

This study considers the interplay between individual load-size selection and recruitment behavior in the leaf-cutting ant Atta cephalotes. Foraging workers anchor themselves on the leaf edge by their hind legs and pivot around them while cutting arcs from leaves. Since workers not only cut leaves but also lay chemical trails to recruit nestmates, we investigated whether there is conflict of motivation affecting the workers' decision either to quickly inform nestmates about a newly discovered food source, or instead to cut full-load leaf fragments, which could delay recruitment. Workers were presented with leaves of privet of three different grades of toughness (measured as leaf density=mass/area) as sources of different quality, and load-size selection and recruitment behavior by harvesting-satiated and harvesting-deprived workers were measured. The following results were obtained. (1) Leaf density affected individual load-size selection: both harvesting-satiated and harvesting-deprived workers were found to cut smaller leaf fragments from the denser leaves. (2) Harvesting-deprived workers cut smaller fragments than harvesting-satiated workers, and therefore saved cutting time. The fragments cut were smaller only during the initial phases of the recruitment process, when information about the discovery needed to be transferred. (3) Harvesting-deprived workers showed higher recruitment rates than harvesting-satiated workers. A considerable number of ants were observed to return to the nest unladen. During the initial phases, the ratio of laden/unladen workers was lower than that for harvesting-satiated workers, and increased with the development of the tograging process. (4) Scout workers confronted with familiar leaves ran back to the nest laying chemical trails without even contacting the leaves. They relied on olfactory cues to start recruiting nestmates, and leaf density played no role in their decisions. (5) When confronted with unfamiliar leaves, on the other hand, they assessed leaf quality by probing bites at the leaf edge, although no actual cuts occurred. In this situation, the resulting recruitment rates depended on physical leaf traits, being higher for the tenderer leaves. (6) Workers foraging on unfamiliar leaves cut smaller fragments than workers cutting familiar leaves, and most of them displayed trail-laying behavior when returning to the nest. The results support the hypothesis of a trade-off between time spent collecting and that invested to recruit nestmates. During the initial phases of exploitation of a newly discovered food source, workers reduced their individual carrying performance in order to return earlier to the colony for further recruitment.     

Factors initiating algal life-form shift from sediment to water

In enclosure and whole-lake experiments, I tested whether life-form shift (recruitment) is a passive process induced by turbulence, a seasonal phenomenon, or a behavior that can be induced by alterations in environmental variables. The number of algal cells recruited from the sediment varied considerably during the experimental period. The most important migrating genera in this study were: Cryptomonas, Dinobryon, Gonyostomum, Gymnodinium, Peridinium, and Synura. An obvious conclusion is that it is not the same factor in each case that causes life-form shift, but that different triggering factors operate in different algal species. Turbulence and temperature were similar in all treatments and therefore did not cause the considerable fluctuations and trends in algal recruitment in the enclosures. This suggests that life0form shift is not a passive process driven by wind and temperature-induced currents. In the enclosure experiment, alterations in the light régime explained a major part (up to 53%) of the variation in recruitment for most genera. For Gymnodinium this was corroborated in the whole-lake experiment, where the depth of the euphotic zone explained 41% of the variation in recruitment. For Cryptomonas, however, 64% of the variation in recruitment was explained by the depth of the oxycline, whereas 52% of the variation in recruitment of Synura were explained by the depths of the euphotic zone and the oxyline. Peridinium pusillum and P. wisconsinense showed low recruitment at high zooplankton abundance and high recruitment at low zoo-plankton abundance in the lake experiment, as well as in the enclosure experiment. Thus, the hypothesis that the presence of grazers can induce shifts in behaviour of some algal groups cannot be rejected.     

Coexistence of coral reef fishes — a lottery for living space

Synopsis Data are summarised from studies of two reef fish communities — pomacentrids territorial on rubble patches, and fishes resident in small isolated colonies of coral. In each case there is evidence that availability of living sites limits numbers of fishes, and that similar species of fish use the same kinds of spaces. Priority of arrival as recruits, rather than subtle differences in requirements or competitive abilities of adults, appears to determine which species holds each site. Faced with a limited and patchy supply of living space, most reef fishes are sedentary as adults, and produce frequent clutches of pelagic larvae over extended breeding seasons In this way they maximise their chances of getting offspring into suitable living sites as such sites appear. It is argued that by adopting this strategy, reef fishes are preadapted for forming inter-specific lotteries for living space if several species with similar requirements occur together. Such lotteries among similar species may be a feature common to many reef fish communities, and may explain the typically high within-site diversity found in them.This paper forms part of the proceedings of a mini-symposium convened at Cornell University, Ithaca, N.Y., 18–19 May 1976, entitled Patterns of Community Structure in Fishes (G. S. Helfman, ed.).     

Habitat selection and juvenile persistence control the distribution of two closely related Caribbean damselfishes

Summary On many Caribbean fringing coral reefs, two closely related and ecologically similar damselfishes, the beaugregory (Stegastes leucostictus Müller and Troschel) and the cocoa damselfish (S. variabilis Castelnau), occupy nonoverlapping vertical distributions. In St. Croix (USVI), beaugregory are very abundant in shallow water back reef habitats (1–2 m depth) while cocoa damselfish are restricted to the base of the forereef (10–15 m depth).In this study, the roles of habitat selection at settlement and juvenile persistence were investigated to determine their influence on this pattern of zonation. Settlement events observed at intervals over a two-year period revealed that habitat selection occurred at settlement and was confined to habitats occupied by adults. In addition, differences in juvenile persistence (due to mortality and/or emigration) were found when species were translocated between depths. Over a period of 100-days, juvenile beaugregory moved from 1 m to 12 m depth suffered four-fold greater losses at the deeper sites than shallow water controls, while translocated cocoa damselfish suffered twice as many losses in shallow water than controls at 12 m depth. Despite these differences in persistence, growth rates of the two species were similar and independent of depth. These results indicate that preferential habitat selection at settlement, perhaps an evolutionary response to differential juvenile mortality, may play a deciding role in determining distributions of ecologically similar species of coral reef fishes.     

Influence of seed size and seed nature on recruitment in the polymorphic harvester ant Messor barbarus

In the polymorphic harvester ant Messor barbarus, the colony modulates response during foraging according to seed characteristics. The easiness of picking up small seeds (oat fragments or canaryseeds) shortens the time lapse between food discovery and the return of scouts to the nest, favouring the onset of recruitment and higher mobilisation rates than bigger seeds (whole oat). However, size is not the only criterion, as canaryseeds are preferred among small ones. A modulation of the trail laid by the first scouts also plays a role in the shaping of harvesting patterns and in the enhancement of recruitment towards small and/or preferred seed. This flexibility extends to the relative participation of the three worker sizes classes, which differs according to seed. Media are the most numerous at the foraging arena and largely responsible for trail-laying. Minor participate in seed harvesting and trail-laying, mostly when they can be efficient in carrying seeds, such as oat fragments. Major scarcely participate in harvesting and trail-laying, but are involved in the exploitation of bigger and/or preferred seed species. When faced with different seed species, M. barbarus seems thus to collectively adopt a “foraging time minimisation” rather than an “energetic gain per item harvested maximisation” strategy. Dynamics of foraging and division of labour between foragers of different sizes are related to both the workers’ ability to carry seed items, and the transfer of information to nestmates by trail-laying, all parameters that vary according to seed characteristics.     

High mobility of the protandrous anemonefish Amphiprion frenatus: nonrandom pair formation in limited shelter space

Protandry in anemonefishes has been attributed to random pair formation, which results from their limited mobility after random recruitment to isolated host anemones. The recruitment and movement of the anemonefish Amphiprion frenatus were investigated in relation to its group structure and the spatial distribution of its host anemone on a coral reef, where it inhabits isolated single hosts and interhost movement is rare. A juvenile tended to be recruited to a host from which a former resident(s) had disappeared, indicating that larval recruitment is not random. After mass bleaching of corals during which many hosts died, a quarter of adults moved between hosts on average 42m in 3 weeks, indicating that their mobility is potentially high. The few migrations under normal conditions were probably due to the low benefit of movements. Even after the movements, a female was much larger than her mate in a host, and a large female tended to pair with a large male. Although body size of females was positively correlated with their host size, that of males was not. It is suggested that the size of a single host does not allow the coexistence of two or more large fish, and the size composition of each pair is affected by the host size. Protandry in the monogamous fish may be attributed to the nonrandom pair formation in the limited space of a host.     

Rodent seed predation promotes differential recruitment among bird-dispersed trees in temperate secondary forests

We investigated the role of seed predation by rodents in the recruitment of the fleshy-fruited trees Taxus baccata, Ilex aquifolium and Crataegus monogyna in temperate secondary forests in NW Spain. We measured the densities of dispersed seeds, early emerged seedlings, established recruits and adults, at four sites over a period of 2 years. Seed predation among species was compared by seed removal experiments and analysis of rodent larder-hoards. The three species differed markedly in local regeneration patterns. The rank order in the seed rain following decreasing seed density was Ilex, Taxus and Crataegus. However, Crataegus established 3.3 times more seedlings than Taxus. For all species, there was a positive linear relationship between the density of emerged seedlings and seed density, suggesting that recruitment was seed- rather than microsite-limited. A consistent pattern of seed selection among species was exerted by rodents, which preferred Taxus and, secondarily, Ilex seeds to Crataegus seeds. Predation ranking was the inverse of that of seed protection against predators, measured as the mass of woody coat per mass unit of the edible fraction. Recruitment potential, evaluated as the ratio of seedlings to seeds, was negatively related to seed predation, with the rank order Crataegus > Ilex > Taxus. The selective early recruitment limitation exerted by predation may have a demographic effect in the long term, as judged by the positive relationship between early seedling emergence and the density of established recruits. By modulating the pre-emptive competition for seed safe sites, rodents may preclude the progressive exclusion of species that produce low numbers of seeds (i.e. Crataegus) by those dominant in seed number (i.e. Ilex, Taxus), or at least foster the evenness for site occupation among seedlings of different species.     

Density dependence at some time and place?

There appears to be widespread acceptance that for a population to persist, some demographic parameter must be density dependent at some place or time. In this paper, we question the veracity and heuristic value of treating this statement as a general principle of ecology. We also point out that some processes that have recently been defined as density dependent are, in fact, not. Taken in its original sense, density dependence implies a change in demographic rates based on biological (generally negative) feedback. Situations exist, however, in which demographic rates change in relation to density without negative biological feedback. For example, per capita recruitment in marine populations will decrease as local population size increases even as absolute numbers of arriving larvae do not change. The failure to separate these density-related processes from true density-dependent processes affects our understanding of population regulation and of the way in which the natural world functions. Furthermore, focusing solely on density-dependent processes and their role in population regulation neglects to address numerous density-independent processes like disturbance and climatic variation that may have important impacts in determining population size. Received: 20 January 1999 / Accepted: 12 January 2000