Fissioning minimizes ranging costs in spider monkeys: a multiple-level approach

The adjustment to deal with intragroup food competition is probably the most plausible explanation of high levels of fission–fusion dynamics. However, studies did not always support expected relations between food availability, ranging costs, and subgroup size. We used several levels of analysis differing in the time and spatial scale in order to investigate this explanation in spider monkeys. In our study, subgroups were larger when food availability was higher across most levels of analyses used. We also found a fine-scale adjustment: compared to the food patch previously visited, spider monkeys traveled to larger patches just after fusions. This was not without an immediate travel cost: the interpatch distance and travel time after a fusion were longer than that before the fusion. This rapid adjustment shows the flexibility that fission–fusion dynamics can offer. Spider monkeys are in large subgroups only when food conditions are favorable, as evidenced by the fact that at all the other time-scale levels larger subgroups did not experience greater ranging costs than smaller subgroups. Our results indicate that on the whole spider monkeys successfully minimize ranging costs by fission and fusion of subgroups.     

Ecological constraints on group size: an analysis of spider monkey and chimpanzee subgroups

The social organization of spider monkeys (Ateles geoffroyi) and chimpanzees (Pan troglodytes) appear remarkably similar. In this paper, field studies of these two species were used to (1) test a model of ecological constraints on animal group size which suggests that group size is a function of travel costs and (2) assess ecological and social factors underlying the social organization of these two species. Spider monkeys were studied over a 6-year period in Santa Rosa National Park, Costa Rica, and chimpanzees were studied for 6 years in Kibale National Park, Uganda. Adults of both species spent their time in small subgroups that frequently changed size and composition. Thus, unlike most primate species, spider monkeys and chimpanzees were not always in a spatially cohesive social group; each individual had the option of associating in subgroups of a different size or composition. Both species relied on ripe fruit from trees that could be depleted through their feeding activity. However, spider monkey food resources tended to occur at higher densities, were more common, less temporally variable, and did not reach the low levels experienced by chimpanzees. Analyses of the relationship between subgroup size and the density and distribution of their food resources suggested that travel costs limit subgroup size. However, these ecological factors did not influence all age/sex classes equally. For example, the number of adult males in a subgroup was a function of food density and travel costs. However, this was not the case for female chimpanzees, suggesting that the benefits of being in a subgroup for females did not exceed the costs, even when ecological conditions appeared to minimize subgroup foraging costs. Therefore, it seems likely that social strategies influenced the relationship between food resource variables and subgroup size.     

To fission or to fusion: effects of community size on wild chimpanzee (<Emphasis Type="Italic">Pan troglodytes verus</Emphasis>) social organisation

To balance advantages and disadvantages of group living, some species have fission-fusion social systems in which members of the same group form frequently changing subgroups. This allows flexible responses of group size to external conditions while at the same time retaining group stability. In chimpanzees, subgroup (party) size and composition depend mainly on the presence of receptive females, food availability and the activity of the party. Here we analyse the extent to which fission-fusion parameters are influenced by changes of demographic variables like community size and composition. Data were collected from a habituated West African chimpanzee community (Taï forest, Côte dIvoire) over 10 years, during which total community size decreased from 51 to 21, and the number of adult males decreased from 9 to 2. Taï chimpanzees are highly gregarious, as they spend more than 80% of their time with unrelated conspecifics. With decreasing community size, party size, party duration and male-female association increased. Neither activity nor the presence of receptive females or feeding competition could explain the observed changes in grouping patterns. Thus, the decrease in community size led to an increase of party cohesion and also enhanced cohesiveness between the sexes, while general sociality remained unchanged. Therefore, our data support the notion that small communities are more cohesive and have a less flexible fission-fusion system.Communicated by D. Watts     

Trade-offs between time, predation risk and life history, and their implications for biogeography: A systems modelling approach with a primate case study

Group sizes are often considered to be the result of a trade-off between predation risk and the costs of feeding competition. We develop a model to explore the interaction between different ecological constraints on group sizes, using a primate (baboons) case study. The model uses climatic correlates of time budgets to predict maximum ecologically tolerable group size, and climatic predictors of predation risk (reflected mainly in predator density and female body mass) to predict minimum tolerable group size for any given habitat. As well as defining the range of sustainable group sizes for a given habitat, the model also allows us to reliably predict our exemplar taxon's biogeographical distribution across Africa. We also explore the life history implications of the model to ask whether baboons form group sizes which maximise survival or fecundity in the classic trade off between these two key life history variables. Our results indicate that, within the range of study sites in our sample, baboons prefer to maximise fecundity. However, the data indicate that in higher predation risk habitats they would switch to maximising survival at the expense of fecundity. We argue that this is due to the fact that interbirth interval and developmental rates have a ceiling that cannot be breached. Thus, while females can shorten interbirth intervals to compensate for increased predation risk, there is a limit to how much these life history variables can be altered, and when this is reached the best strategy is to maximise survivorship.     

Movements of vervets (Cercopithecus aethiops) and patas monkeys (Erythrocebus patas) as estimators of food resource size, density, and distribution

The effect of food resources on behavior has been difficult to measure. Here we use animals themselves to describe “effective” food abundance and distribution by comparing, relative to where individuals stopped to eat, movements of (1) adult females living in a small group of vervet monkeys (Cercopithecus aethiops) with those living in a large group and (2) vervets and patas monkeys (Erythrocebus patas). Although females in the large vervet group travelled farther and stopped to eat more often than females in the small vervet group, these differences resulted from foraging in Acacia drepanolobium habitat. In A. xanthophloea habitat, females in the large group travelled less far, travelled shorter distances between foods, and stopped as often as females in the small group. Greater foraging costs of females in larger vervet groups may be offset by access to home ranges of better quality. Compared to patas, vervets travelled shorter distances, moved shorter distances between food sites, stopped less often, and had longer feeding bouts, suggesting that foods of vervets are denser and larger, overall, than foods of patas. When vervets foraged in A. drepanolobium habitat, also the habitat of patas, their foraging behavior became more like that of patas. Vervets travelled farther, stopped more often, and spent less time at food sites in A. drepanolobium habitat than in A. xanthophloea habitat, suggesting that foods are smaller and less usurpable in A. drepanolobium habitat. Distance between foods, a component of food distribution, did not increase, however. The critical variable underlying usurpability of foods may be food site depletion time, a temporal measure. Received: 14 March 1997 / Accepted after revision: 19 October 1997     

Scale-dependent variation in coral community similarity across sites, islands, and island groups

Community similarity is the proportion of species richness in a region that is shared on average among communities within that region. The slope of local richness (alpha diversity) regressed on regional richness (gamma diversity) can serve as an index of community similarity across regions with different regional richness. We examined community similarity in corals at three spatial scales (among transects at a site, sites on an island, and islands within an island group) across a 10 000-km longitudinal diversity gradient in the west-central Pacific Ocean. When alpha diversity was regressed on gamma diversity, the slopes, and thus community similarity, increased with scale (0.085, 0.261, and 0.407, respectively) because a greater proportion of gamma diversity was subsumed within alpha diversity as scale increased. Using standard randomization methods, we also examined how community similarity differed between observed and randomized assemblages and how this difference was affected by spatial separation of species within habitat types and specialization of species to three habitat types (reef flats, crests, and slopes). If spatial separation within habitat types and/or habitat specialization (i.e., underdispersion) occurs, fewer species are shared among assemblages than the random expectation. When the locations of individual coral colonies were randomized within and among habitat types, community similarity was 46-47% higher than that for observed assemblages at all three scales. We predicted that spatial separation of coral species within habitat types should increase with scale due to dispersal/extinction dynamics in this insular system, but that specialization of species to different habitat types should not change because habitat differences do not change with scale. However, neither habitat specialization nor spatial separation within habitat types differed among scales. At the two larger scales, each accounted for 22-24% of the difference in community similarity between observed and randomized assemblages. At the smallest scale (transect-site), neither spatial separation within habitat types nor habitat specialization had significant effects on community similarity, probably due to the small size of transect samples. The results suggest that coral species can disperse among islands in an island group as easily as they can among sites on an island over time scales that are relevant to their establishment and persistence on reefs.     

Organization of Plethodon salamander communities: guild-based community assembly

A long-standing goal in evolutionary ecology is to determine whether the organization of communities is reflective of underlying deterministic processes. In this study, I examined patterns of species co-occurrence among eastern Plethodon salamanders and determined whether they were consistent with predictions from a guild model of competition-based community assembly. Using a database of 45 species and 4540 geographic sites, I found that patterns of co-occurrence were significantly nonrandom at both a regional and continental scale, and species of different size guilds were distributed more evenly in sites than was expected by chance. Sites with the highest species richness had consistent patterns of community composition, and with few exceptions, the same five species were present at all sites. Taken together, these results imply that larger Plethodon communities are assembled from simpler communities in a manner consistent with what is predicted through competitive mechanisms and suggest that stable species combinations are possible to achieve at various levels of species richness. These results also provide strong evidence consistent with the hypothesis that competitive-based community assembly is a general phenomenon in Plethodon and that interspecific competition is prevalent among the eastern species of this group.     

The impact of grazing on spider communities in a mesophytic calcareous dune grassland

During 1994–1995 and 1997–1998 spiders were sampled with pitfall traps in a botanically rich, mesophytic, calcareous dune grassland in Belgium. As a consequence of intensive cattle grazing, vegetation variation in a large part of the area had diminished. The study area was also patchily grazed by rabbits. Community analysis with TWINSPAN revealed five distinct spider communities. Ecological differentiation was best explained by combination of the habitat variables: distance from grazed or non-grazed vegetation,Rosa pimpinellifolia cover and grass cover in both summer and winter. Species diversity was highest in the border zone between the cattle-grazed and non cattle-grazed sites. Correlation of the most abundant spider species with the vegetation determinants explains the ecological differentiation between the spider communities. Species were classified into seven major groups that reflect the species’ habitat preferences. The group showing clear association with non cattle-grazed, tall vegetation consists of common species. Characteristic species for the intensively cattle-grazed sites are common aeronauts and rare species such asWalckenaeria stylifrons, Mastigusa arietina, Ceratinopsis romana andPardosa monticola. The latter are shown to be dependent on ungrazed vegetation for juvenile development and overwintering. Intensive grazing results in homogeneous short vegetation, which can only be colonized by ‘open ground’ species with a well-developed dispersal capacity, or by species which are not dependent on litter-rich situations for juvenile development. An extensive cattle grazing regime results in a patchy mosaic grassland where, in addition to the above mentioned groups of species, other species survive by migrating between the buffered litter rich ungrazed vegetation and the short vegetation. Additionally, some typical and rare species prefer the transition zone between the grazed and the ungrazed vegetation because they are associated with specific habitat structures or inhabiting ant-species.     

Geographic variation in social organization of Galápagos mockingbirds: ecological correlates of group territoriality and cooperative breeding

Summary To investigate ecological influences on cooperative social organization, I studied the four allopatric species of mockingbirds (Nesomimus spp.) endemic to the Galápagos archipelago on four islands. On three small, low and arid islands (Genovesa, Champion and Española), mockingbird territories filled all terrestrial habitat, mean group size varied from 4.5 to 14.2 adults, maximum group size ranged from seven to 24 birds, and 70–100% of groups contained more than two birds. San Cristóbal is larger and higher, and it supports a broader range of habitats. At one highland and two coastal sites on this island, mockingbirds did not hold territories in all available habitats, group size averaged 2.2 adults, only 25% of groups were larger than two, and none included more than three adults. Adults dispersed into vacant habitat to establish new territories only on San Cristóbal. Helping behavior has not yet been observed on San Cristóbal, but it occurs on the other three islands. These results support the hypothesis that social groups and cooperative breeding are maintained where limited availability of preferred habitat constrains dispersal. The mechanism relaxing habitat saturation on San Cristóbal, however, remains undetermined. Predation by introduced rats and cats may reduce survival and indirectly reduce group size; these predators are absent from Genovesa, Champion and Española. Differences in food supplies could also affect interand intra-island variation in population density. Variation in social organization among arid coastal sites on the four islands, and similarity between climatically different sites on San Cristóbal, suggest that climatic conditions are less important as determinants of dispersal and breeding. Skews in adult sex ratios also fail to account for inter-island variation in sociality. Although they live in a climatically variable environment, territorial behavior and the physical limits of suitable habitat have an overriding influence on cooperative social organization in Galápagos mockingbirds.     

Quantifying relationships between bird and butterfly community shifts and environmental change.

Quantifying the manner in which ecological communities respond during a time of decreasing precipitation is a first step in understanding how they will respond to longer-term climate change. Here we coupled analysis of interannual variability in remotely sensed data with analyses of bird and butterfly community changes in montane meadow communities of the Greater Yellowstone Ecosystem. Landsat satellite imagery was used to classify these meadows into six types along a hydrological gradient. The northern portion of the ecosystem, or Gallatin region, has smaller mean patch sizes separated by ridges of mountains, whereas the southern portion of the ecosystem, or Teton region, has much larger patches within the Jackson Hole valley. Both support a similar suite of butterfly and bird species. The Gallatin region showed more overall among-year variation in the normalized difference vegetation index (NDVI) when meadow types were pooled within regions, perhaps because the patch sizes are smaller on average. Bird and butterfly communities showed significant relationships relative to meadow type and NDVI. We identified several key species that are tightly associated with specific meadow types along the hydrological gradient. Comparing taxonomic groups, fewer birds showed specific habitat affinities than butterflies, perhaps because birds are responding to differences in habitat structure among meadow types and using the landscape at a coarser scale than the butterflies. Comparing regions, the Teton region showed higher predictability of community assemblages as compared to the Gallatin region. The Gallatin region exhibited more significant temporal trends with respect to butterflies. Butterfly communities in wet meadows showed a distinctive shift along the hydrological gradient during a drought period (1997-2000). These results imply that the larger Teton meadows will show more predictable (i.e., static) species-habitat associations over the long term, but that the smaller Gallatin meadows may be an area that will exhibit the effects of global climate change faster.     

Habitat fragmentation and effects of herbivore (howler monkey) abundances on bird species richness

Habitat fragmentation can alter herbivore abundances, potentially causing changes in the plant community that can propagate through the food web and eventually influence other important taxonomic groups such as birds. Here we test the relationship between the density of red howler monkeys (Alouatta seniculus) and bird species richness on a large set of recently isolated land-bridge islands in Lago Guri, Venezuela (n = 29 islands). Several of these islands host relict populations of howler monkeys at densities up to more than 30 times greater than those on the mainland. These "hyperabundant" herbivores previously have been shown to have a strong positive influence on aboveground plant productivity. We predicted that this should lead to a positive, indirect effect of howler monkey density on bird species richness. After accounting for passive sampling (the tendency for species richness to be positively associated with island area, regardless of differences in habitat quality) we found a significant positive correlation between howler monkey density and bird species richness. A path analysis incorporating data on tree growth rates from a subset of islands (n = 9) supported the hypothesis that the effect of howler monkeys on the resident bird communities is indirect and is mediated through changes in plant productivity and habitat quality. These results highlight the potential for disparate taxonomic groups to be related through indirect interactions and trophic cascades.     

Correlations of food distribution and patch size with agonistic interactions in female vervets (Chlorocebus aethiops) and patas monkeys (Erythrocebus patas) living in simple habitats

Food distribution is hypothesized to be important in determining the nature of female relationships within social groups of primates. When food limits female reproductive success, spatially clumped foods are expected to produce strong, linear dominance hierarchies within groups, whereas more spatially dispersed foods are expected to produce weaker or non-existent dominance hierarchies. The association between food distribution and competitive relationships presumably occurs because clumped foods are usurpable but dispersed foods are not. We examined the spatial distribution of food patches (trees) and patch size relative to feeding behavior and agonistic interactions in vervets and patas monkeys, two closely related and sympatric species that nonetheless differ in the strength of the female dominance hierarchy. Food patches of both patas monkeys and vervets were small in size and randomly distributed in Acacia drepanolobium habitat. In contrast, in A. xanthophloea woodland, the habitat type that was exclusively used by vervets, food patches were larger and more spatially clumped. These similarities and differences between and within species were correlated with similarities and differences in the strength and linearity of their dominance hierarchies. Patas monkeys and vervets in A. drepanolobium habitat had dominance hierarchies that were weakly defined because there were relatively few agonistic interactions between females. By contrast, in A. xanthophloea habitat, vervets had a stronger, linear dominance hierarchy characterized by a higher rate of agonistic interactions over food. The covariation of agonistic interactions with patch size is discussed in relation to depletion time, another characteristic that may covary with food distribution, and resource renewal rate, an important determinant of agonistic interactions in insectivorous birds, fishes, insects, and mammals. Received: 18 February 2000 / Revised: 5 September 2000 / Accepted: 26 September 2000     

Flows, droughts, and aliens: factors affecting the fish assemblage in a Sierra Nevada, California, stream

The fishes of Martis Creek, in the Sierra Nevada of California (USA), were sampled at four sites annually over 30 years, 1979-2008. This long-term data set was used to examine (1) the persistence and stability of the Martis Creek fish assemblage in the face of environmental stochasticity; (2) whether native and alien fishes responded differently to a natural hydrologic regime (e.g., timing and magnitude of high and low flows); and (3) the importance of various hydrologic and physical habitat variables in explaining the abundances of native and alien fish species through time. Our results showed that fish assemblages were persistent at all sample sites, but individual species exhibited marked interannual variability in density, biomass, and relative abundance. The density and biomass of native fishes generally declined over the period of study, whereas most alien species showed no significant long-term trends. Only alien rainbow trout increased in both density and biomass at all sites over time. Redundancy analysis identified three hydrologic variables (annual 7-day minimum discharge, maximum winter discharge, and number of distinct winter floods) and two habitat variables (percentage of pool habitat and percentage of gravel substrate) that each explained a significant portion of the annual variation in fish assemblage structure. For alien taxa, their proportional contribution to the total fish assemblage was inversely related to mean annual streamflow, one-day maximum discharge in both winter and spring, and the frequency of springtime floods. Results of this study highlight the need for continuous annual monitoring of streams with highly variable flow regimes to evaluate shifts in fish community structure. Apparent successes or failures in stream management may appear differently depending on the time series of available data.     

Defoliation synchronizes aboveground growth of co-occurring C4 grass species

The aboveground net primary productivity (ANPP) of grass communities in grasslands and savannas is primarily determined by precipitation quantity. Recent research, motivated by predictions of changes in the distribution of rainfall events by global climate change models, indicates that ANPP may be affected by rainfall distribution as much as by annual totals. Grazing and community composition are also known to affect grassland ANPP. The manner in which interactions between rainfall distribution, grazing, and community composition affect the relationship between precipitation and ANPP represents a critical knowledge gap. The effects of community composition and grazing on aboveground growth responses to intraseasonal variation in water availability were investigated at seven grassland sites with a nonselective clipping experiment. The aboveground growth of the dominant C4 species at each site was measured at regular intervals for 2-3 growing seasons in the presence or absence of regular defoliation. In the absence of defoliation, there was a general lack of synchrony of intraseasonal growth among co-occurring species. Variation in growth rates was high and was only partially explained by variation in rainfall. Regular defoliation increased growth synchrony at all sites, but changes in growth responses to rainfall varied between sites. These results suggest that community composition will be important in determining ANPP-precipitation relationships under conditions of altered rainfall distribution. However this effect appears to be a result of species responding differently to soil water or other resources rather than to rainfall per se. Grazing may override the effects of community composition by reducing differences in growth patterns between species and has the potential to weaken precipitation controls on ANPP.     

Climate, habitat, and species interactions at different scales determine the structure of a Neotropical bat community

Climate, habitat, and species interactions are factors that control community properties (e.g., species richness, abundance) across various spatial scales. Usually, researchers study how a few properties are affected by one factor in isolation and at one scale. Hence, there are few multi-scale studies testing how multiple controlling factors simultaneously affect community properties at different scales. We ask whether climate, habitat structure, or insect resources at each of three spatial scales explains most of the variation in six community properties and which theory best explains the distribution of selected community properties across a rainfall gradient. We studied a Neotropical insectivorous bat ensemble in the Isthmus of Panama with acoustic monitoring techniques. Using climatological data, habitat surveys, and insect captures in a hierarchical sampling design we determined how much variation of the community properties was explained by the three factors employing two approaches for variance partitioning. Our results revealed that most of the variation in species richness, total abundance, and feeding activity occurred at the smallest spatial scale and was explained by habitat structure. In contrast, climate at large scales explained most of the variation in individual species' abundances. Although each species had an idiosyncratic response to the gradient, species richness peaked at intermediate levels of precipitation, whereas total abundance was very similar across sites, suggesting density compensation. All community properties responded in a different manner to the factor and scale under consideration.     

Relative size of Aristotle's lantern in Echinometra mathaei occuring at different densities

Like species of sea urchins in Zanzibar and Oregon (USA), Echinometra mathaei (de Blainville) at Rottnest Island, Western Australia, displays variation in the size of Aristotle's lantern relative to the maximum diameter of the test. This variation was associated with local variations in density of urchins at each of two sites in each of two years (1980 and 1981); this association with density was consistent with the proposal that relatively larger lanterns are a response to decreased food availability. Furthermore, variation of relative lantern size associated with local density was similar in magnitude to the variation displayed between sites and between years. This temporal variation demonstrated the plasticity of the relative lantern size over periods as short as 12 mo. Further experimental studies are required before relative length of lanterns can be used as estimates of food availability.     

The impact of grazing on spider communities in a mesophytic calcareous dune grassland

During 1994–1995 and 1997–1998 spiders were sampled with pitfall traps in a botanically rich, mesophytic, calcareous dune grassland in Belgium. As a consequence of intensive cattle grazing, vegetation variation in a large part of the area had diminished. The study area was also patchily grazed by rabbits. Community analysis with TWINSPAN revealed five distinct spider communities. Ecological differentiation was best explained by combination of the habitat variables: distance from grazed or non-grazed vegetation,Rosa pimpinellifolia cover and grass cover in both summer and winter. Species diversity was highest in the border zone between the cattle-grazed and non cattle-grazed sites. Correlation of the most abundant spider species with the vegetation determinants explains the ecological differentiation between the spider communities. Species were classified into seven major groups that reflect the species’ habitat preferences. The group showing clear association with non cattle-grazed, tall vegetation consists of common species. Characteristic species for the intensively cattle-grazed sites are common aeronauts and rare species such asWalckenaeria stylifrons, Mastigusa arietina, Ceratinopsis romana andPardosa monticola. The latter are shown to be dependent on ungrazed vegetation for juvenile development and overwintering. Intensive grazing results in homogeneous short vegetation, which can only be colonized by ‘open ground’ species with a well-developed dispersal capacity, or by species which are not dependent on litter-rich situations for juvenile development. An extensive cattle grazing regime results in a patchy mosaic grassland where, in addition to the above mentioned groups of species, other species survive by migrating between the buffered litter rich ungrazed vegetation and the short vegetation. Additionally, some typical and rare species prefer the transition zone between the grazed and the ungrazed vegetation because they are associated with specific habitat structures or inhabiting ant-species. Nomenclature: Roberts (1987, 1995) forAraneae; van der Meijden et al. (1990) for vascular plants; Corly et al. (1981) for bryophytes; Schaminée et al. (1996) for vegetation associations.     

Size-mediated non-trophic interactions and stochastic predation drive assembly and dynamics in a seabird community

Theoretical and empirical evidence suggests that body size is a major life-history trait impacting on the structure and functioning of complex food webs. However, long-term analyses of size-dependent interactions within simpler network modules, for instance, competitive guilds, are scant. Here, we model the assembly dynamics of the largest breeding seabird community in the Mediterranean basin during the last 30 years. This unique data set allowed us to test, through a "natural experiment," whether body size drove the assembly and dynamics of an ecological guild growing from very low numbers after habitat protection. Although environmental stochasticity accounted for most of community variability, the population variance explained by interspecific interactions, albeit small, decreased sharply with increasing body size. Since we found a demographic gradient along a body size continuum, in which population density and stability increase with increasing body size, the numerical effects of interspecific interactions were proportionally higher on smaller species than on larger ones. Moreover, we found that the per capita interaction coefficients were larger the higher the size ratio among competing species, but only for the set of interactions in which the species exerting the effect was greater. This provides empirical evidence for long-term asymmetric interspecific competition, which ultimately prompted the local extinction of two small species during the study period. During the assembly process stochastic predation by generalist carnivores further triggered community reorganizations and global decays in population synchrony, which disrupted the pattern of interspecific interactions. These results suggest that the major patterns detected in complex food webs can hold as well for simpler sub-modules of these networks involving non-trophic interactions, and highlight the shifting ecological processes impacting on assembling vs. asymptotic communities.     

Priority effects and species sorting in a long paleoecological record of repeated community assembly through time

We studied the relative roles of environmental species sorting and priority effects in the assembly of ecological communities on long time scales, by analyzing community turnover of water fleas (Daphnia) in response to strong and recurrent environmental change in a fluctuating tropical lake. During the past 1800 years, Lake Naivasha (Kenya) repeatedly fluctuated between a small saline pond habitat during lowstands and a large freshwater lake habitat during highstands. Starting from a paleoecological reconstruction, we estimated the role of priority effects in Daphnia community assembly across 16 of these habitat turnovers and compared this with the response of the community to reconstructed changes in three environmental variables important for species sorting. Our results indicate that the best predictor of Daphnia community composition during highstands was the community composition just prior to the transition from lowstands to highstands. This reflects a long-lasting priority effect of late lowstand communities on highstand communities, arising when remnant lowstand populations fill newly available ecological space in the rapidly expanding lake habitat. Species sorting and priority effects had a comparable but relatively small influence on community composition during the lowstands. Moreover, these priority effects decayed rapidly with time as Daphnia communities responded to environmental change, in contrast with the highstand communities where priority effects lasted for several decades.     

Species richness patterns along environmental gradients in island land molluscan fauna

The relationship between species richness and environmental variables may change depending on habitat structure, dispersal ability, species mixing, and community adaptation to the environment. It is crucial to know how these factors regulate the environment-diversity relationship. The land molluscan fauna of the Ogasawara Islands in the West Pacific is an excellent model system to address this question because of the high species endemicity (> 90%), small area, and simple habitat structure of the islands. I examined relationships among indigenous species composition, richness, and habitat condition, and especially productivity and forest moisture on the island of Anijima. Two major communities of snails could be distinguished by detrended correspondence analysis (DCA): one group dominated in a moist habitat with high productivity, and the other group dominated in a dry habitat with low productivity. However, species richness became highest at the intermediate condition between the habitats in which the two snail communities were dominant, so that species richness showed a hump-shaped relationship with moisture and productivity. In contrast, the species richness of the snail community in the moist habitat showed a monotonically positive correlation, and that in the dry habitat showed a monotonically negative correlation with moisture and productivity. Thus, the greater species richness in intermediate moisture and productivity resulted from the ecotone effect or community overlap at the transitional areas, where faunas with different ecologies can meet in a single site. These findings suggest that hump-shaped productivity-diversity relationships in land Mollusca would reflect the ecotone effect as a result of the mixing of species adapted to either fertile habitats or sterile habitats.