Contagious seed dispersal beneath heterospecific fruiting trees and its consequences

An hypothesized advantage of seed dispersal is avoidance of high per capita mortality (i.e. density-dependent mortality) associated with dense populations of seeds and seedlings beneath parent trees. This hypothesis, inherent in nearly all seed dispersal studies, assumes that density effects are species-specific. Yet because many tree species exhibit overlapping fruiting phenologies and share dispersers, seeds may be deposited preferentially under synchronously fruiting heterospecific trees, another location where they may be particularly vulnerable to mortality, in this case by generalist seed predators. We demonstrate that frugivores disperse higher densities of Cornus florida seeds under fruiting (female) Ilex opaca trees than under non-fruiting (male) Ilex trees in temperate hardwood forest settings in South Carolina, USA. To determine if density of Cornus and/or Ilex seeds influences survivorship of dispersed Cornus seeds, we followed the fates of experimentally dispersed Cornus seeds in neighborhoods of differing, manipulated background densities of Cornus and Ilex seeds. We found that the probability of predation on dispersed Cornus seeds was a function of both Cornus and Ilex background seed densities. Higher densities of Ilex seeds negatively affected Cornus seed survivorship, and this was particularly evident as background densities of dispersed Cornus seeds increased. These results illustrate the importance of viewing seed dispersal and predation in a community context, as the pattern and intensity of density-dependent mortality may not be solely a function of conspecific densities.     

Active dispersal is differentially affected by inter‐ and intraspecific competition in closely related nematode species

Competition is one of the main drivers of dispersal, which can be an important mechanism to achieve permanent or temporal coexistence of multiple species. This coexistence can be achieved by a dispersal‐competition tradeoff, spatial store effects or neutral dynamics. Here we test the effect of inter‐ and intraspecific competition on dispersal of four species of the marine nematode species complex Litoditis marina. A previous study in closed microcosms without a possibility for dispersal had demonstrated pronounced interspecific competition, leading to the exclusion of one species. We now investigated whether 1) the dispersal is affected by interspecific interactions, by intraspecific competition (density) or by food availability, 2) the dispersal dynamics influence assemblage composition and can lead to co‐occurrence of the species, and 3) the abiotic environment (here salinity) can affect these dynamics. We show that density is the main driver for dispersal in two of the four species. Dispersal of a third species always started at the same time irrespective of density, whereas in the fourth species interspecific interactions accelerated dispersal. Remarkably, this fourth species was not a strong competitor, suggesting that a dispersal–competition tradeoff does not explain the observed coexistence. Salinity did not alter the timing of dispersal when interspecific interactions were present but did affect assemblage composition. Consequently, spatial store effects may influence coexistence. All four species co‐occurred in fairly stable abundances throughout the present experiment indicating the importance of species specific dispersal strategies for coexistence. Co‐occurrence can be facilitated because competition is postponed or avoided by dispersal. Neutral dynamics also played a role as intra‐ and interspecific competition were of similar importance in three of the four species. We conclude that dispersal is a driver of the coexistence of closely related nematode species, and that population density and interspecific interactions shape these dynamics.     

Spatially correlated recruitment of a marine predator and its prey shapes the large-scale pattern of density-dependent prey mortality

Patterns of predator dispersal can be critical to the dynamics of prey metapopulations. In marine systems, oceanic currents may shape the dispersal of planktonic larvae of both predators and prey, producing spatial correlations in the recruitment of both species and distinctive geographic patterns of prey mortality. I examined the potential for this phenomenon in two fishes, a wrasse and its grouper predator, at a Caribbean island where the near-shore oceanographic regime produces a temporally consistent spatial pattern of fish recruitment. I found that recruitment and adult abundance of groupers were spatially correlated with recruitment of wrasse prey. Furthermore, the local abundance of predators strongly affected the nature of density-dependent prey mortality. At sites with few predators, wrasse mortality was inversely density-dependent, while mortality was positively density-dependent at sites with higher predator densities. This phenomenon could be important to the dynamics of any metacommunity in which physical forces produce correlated dispersal.     

Coexisting cryptic species of the Litoditis marina complex (Nematoda) show differential resource use and have distinct microbiomes with high intraspecific variability

Differences in resource use or in tolerances to abiotic conditions are often invoked as potential mechanisms underlying the sympatric distribution of cryptic species. Additionally, the microbiome can provide physiological adaptations of the host to environmental conditions. We determined the intra‐ and interspecific variability of the microbiomes of three cryptic nematode species of the Litoditis marina species complex that co‐occur, but show differences in abiotic tolerances. Roche 454 pyrosequencing of the microbial 16S rRNA gene revealed distinct bacterial communities characterized by a substantial diversity (85–513 OTUs) and many rare OTUs. The core microbiome of each species contained only very few OTUs (2–6), and four OTUs were identified as potentially generating tolerance to abiotic conditions. A controlled experiment in which nematodes from two cryptic species (Pm1 and Pm3) were fed with either an E. coli suspension or a bacterial mix was performed, and the 16S rRNA gene was sequenced using the MiSeq technology. OTU richness was 10‐fold higher compared to the 454 data set and ranged between 1118 and 7864. This experiment confirmed the existence of species‐specific microbiomes, a core microbiome with few OTUs, and high interindividual variability. The offered food source affected the bacterial community and illustrated different feeding behaviour between the cryptic species, with Pm3 exhibiting a higher degree of selective feeding than Pm1. Morphologically similar species belonging to the same feeding guild (bacterivores) can thus have substantial differences in their associated microbiomes and feeding strategy, which in turn may have important ramifications for biodiversity–ecosystem functioning relationships.     

Untangling the mechanisms of cryptic species coexistence in a nematode community through individual-based modelling

Cryptic species are morphologically identical but genetically distinct, and are prominent across numerous phyla. The coexistence of such closely related species on local scales would seem to run counter to traditional coexistence and competition theory; it has been hypothesized as a consequence of differences in their resource use or tolerances to environmental conditions. We developed an individual-based model of a community of three cryptic Litoditis marina (nematode) species, to understand how individual-level interspecific and intraspecific interactions might explain the coexistence of these closely related species. The model incorporates individuals' reproduction, competition, dispersal and resource use. Data characterizing the cryptic species (growth rates, dispersal ability, competitive interactions and responses to changing environmental conditions) were obtained from laboratory experiments involving both mono- and multispecific nematode cultures, and are used to parameterize the model. Simulation studies are used to investigate which individual-level mechanisms of dispersal and interaction lead to the characteristic population-level patterns observed experimentally. Our results highlight the key role of intraspecific competition in mediating dispersal and therefore co-occurrence of the cryptic species. The differences in dispersal also influence the response of the cryptic species to competition, a combination of factors that provides an explanation for their co-occurrence. These results provide insights into how changes in individual-level processes can be amplified to affect population-level co-occurrence.     

Phylogeography of the Rhabditis (Pellioditis) marina species complex: evidence for long-distance dispersal, and for range expansions and restricted gene flow in the northeast Atlantic

Pinpointing processes that structure the geographical distribution of genetic diversity of marine species and lead to speciation is challenging because of the lack of obvious dispersal barriers and the likelihood of substantial (passive) dispersal in oceans. In addition, cryptic radiations with sympatric distributions abound in marine species, challenging the allopatric speciation mechanism. Here, we present a phylogeographical study of the marine nematode species complex Rhabditis ( Pellioditis ) marina to investigate processes shaping genetic structure and speciation. Rhabditis ( P .) marina lives on decaying macroalgae in the intertidal, and may therefore disperse over considerable distances. Rhabditis ( P .) marina consists of several cryptic species sympatrically distributed at a local scale. Genetic variation in the COI gene was screened in 1362 specimens from 45 locations around the world. Two nuclear DNA genes (ITS and D2D3) were sequenced to infer phylogenetic species. We found evidence for ten sympatrically distributed cryptic species, seven of which show a strong genetic structuring. A historical signature showed evidence for restricted gene flow with occasional long-distance dispersal and range expansions pre-dating the last glacial maximum. Our data also point to a genetic break around the British Isles and a contact zone in the Southern Bight of the North Sea. We provide evidence for the transoceanic distribution of at least one cryptic species (PmIII) and discuss the dispersal capacity of marine nematodes. The allopatric distribution of some intraspecific phylogroups and of closely related cryptic species points to the potential for allopatric speciation in R. ( P .) marina .     

Impacts of grazing soil fauna on decomposer fungi are species-specific and density-dependent

The grazing impacts of different densities of woodlice, collembola and millipedes on the foraging and distribution of two saprotrophic cord-forming basidiomycetes were investigated in soil microcosms. Effects of all three invertebrate species were density-dependent, with larger populations limiting mycelial development to a greater extent. Impacts were, however, species-specific; grazing pressures exerted by low-density woodlouse populations outweighed those of high-density millipede or collembola populations. The varying abilities of soil invertebrates to influence mycelial foraging and distribution indicate that invertebrate species composition and diversity may be key factors regulating saprotrophic basidiomycete functioning in woodland soil.     

The natural history of a fugitive prairie plant (Mirabilis hirsuta (Pursh) MacM.)

Summary Some perennial fugitive plants that colonize badger disturbances in xeric prairies have a limited dispersal capacity, and consequently propagules are dispersed over a small area. I hypothesized that high density-dependent mortality might occur early in the life history of such species, and thus increased survival might occur in subsequent age classes because intraspecific competition would be reduced. These hypotheses were tested using natural and experimental cohorts of Mirabilis hirsuta (Pursh) MacM. From these data and field observations, inferences were obtained concerning selective forces operating upon life history characteristics of this species.The distance between individuals of M. hirsuta increases in successive age classes; the greatest decrease in density occurs between the propagule and seedling age classes. Mortality of propagules due to predation by ants and mice was density-dependent. Predation rates were highest at high propagule densities and predation upon propagules located on badger disturbances was higher than the mortality of propagules at similar densities in undisturbed prairie. The results of mortality in the propagule age class are seedlings present only at low densities and located away from parent plants. Seedlings survive to maturity only if they are located on badger disturbances; this species apparently can not successfully compete with plants present in undistrubed prairie. On badger disturbances seedlings present at low densities have much higher survival (roughly 50%) to maturity than do seedlings present at high densities (essentially zero). Thus, if high densities of propagules occur on a disturbance, predation upon propagules results, indirectly, in increased survival of seedlings to maturity. Such predation potentially could have important effects upon interspecific competition of M. hirsuta with other fugitives also colonizing badger disturbances.Reproductive success of M. hirsuta on the Cayler Prairie Preserve is contingent upon successful colonization of disturbance sites. It would appear that selection has operated upon the life history characteristics to favor both successful immigration onto new sites and establishment of seedlings on those sites. Relatively few, but large propagules are produced annually over a long adult life span. While large propagules enhance seedling establishment on xeric sites, production of few propagules annually for a number of years increases the likelihood of immigration onto sites that are variable in the time of appearance within the dispersal range of the plant.     

The evolution of density-dependent dispersal

Despite a large body of empirical evidence suggesting that the dispersal rates of many species depend on population density, most metapopulation models assume a density-independent rate of dispersal. Similarly, studies investigating the evolution of dispersal have concentrated almost exclusively on density-independent rates of dispersal. We develop a model that allows density-dependent dispersal strategies to evolve. Our results demonstrate that a density-dependent dispersal strategy almost always evolves and that the form of the relationship depends on reproductive rate, type of competition, size of subpopulation equilibrium densities and cost of dispersal. We suggest that future metapopulation models should account for density-dependent dispersal     

Density-dependent effects of larval dispersal mediated by host plant quality on populations of an invasive insect

The success of invasive species is often thought to be due to release from natural enemies. This hypothesis assumes that species are regulated by top-down forces in their native range and are likely to be regulated by bottom-up forces in the invasive range. Neither of these assumptions has been consistently supported with insects, a group which includes many destructive invasive species. Winter moth (Operophtera brumata) is an invasive defoliator in North America that appears to be regulated by larval mortality. To assess whether regulation was caused by top-down or bottom-up forces, we sought to identify the main causes of larval mortality. We used observational and manipulative field and laboratory studies to demonstrate that larval mortality due to predation, parasitism, and disease were minimal. We measured the response of larval dispersal in the field to multiple aspects of foliar quality, including total phenolics, pH 10 oxidized phenolics, trichome density, total nitrogen, total carbon, and carbon–nitrogen ratio. Tree-level declines in density were driven by density-dependent dispersal of early instars. Late instar larvae dispersed at increased rates from previously damaged as compared to undamaged foliage, and in 2015 field larval dispersal rates were related to proportion of oxidative phenolics. We conclude that larval dispersal is the dominant source of density-dependent larval mortality, may be mediated by induced changes in foliar quality, and likely regulates population densities in New England. These findings suggest that winter moth population densities in New England are regulated by bottom-up forces, aligning with the natural enemy release hypothesis.     

Spatial variation and density-dependent dispersal in competitive coexistence

It is well known that dispersal from localities favourable to a species' growth and reproduction (sources) can prevent competitive exclusion in unfavourable localities (sinks). What is perhaps less well known is that too much emigration can undermine the viability of sources and cause regional competitive exclusion. Here, I investigate two biological mechanisms that reduce the cost of dispersal to source communities. The first involves increasing the spatial variation in the strength of competition such that sources can withstand high rates of emigration; the second involves reducing emigration from sources via density-dependent dispersal. I compare how different forms of spatial variation and modes of dispersal influence source viability, and hence source-sink coexistence, under dominance and pre-emptive competition. A key finding is that, while spatial variation substantially reduces dispersal costs under both types of competition, density-dependent dispersal does so only under dominance competition. For instance, when spatial variation in the strength of competition is high, coexistence is possible (regardless of the type of competition) even when sources experience high emigration rates; when spatial variation is low, coexistence is restricted even under low emigration rates. Under dominance competition, density-dependent dispersal has a strong effect on coexistence. For instance, when the emigration rate increases with density at an accelerating rate (Type III density-dependent dispersal), coexistence is possible even when spatial variation is quite low; when the emigration rate increases with density at a decelerating rate (Type II density-dependent dispersal), coexistence is restricted even when spatial variation is quite high. Under pre-emptive competition, density-dependent dispersal has only a marginal effect on coexistence. Thus, the diversity-reducing effects of high dispersal rates persist under pre-emptive competition even when dispersal is density dependent, but can be significantly mitigated under dominance competition if density-dependent dispersal is Type III rather than Type II. These results lead to testable predictions about source-sink coexistence under different regimes of competition, spatial variation and dispersal. They identify situations in which density-independent dispersal provides a reasonable approximation to species' dispersal patterns, and those under which consideration of density-dependent dispersal is crucial to predicting long-term coexistence.     

Density-dependent dispersal complicates spatial synchrony in tri-trophic food chains

Spatial synchrony can increase extinction risk and undermines metapopulation persistence. Both dispersal and biotic interactions can strongly affect spatial synchrony. Here, we explore the spatial synchrony of a tri-trophic food chain in two patches connected by density-dependent dispersal, namely the strategies of prey evasion (PE) and predator pursuit (PP). The dynamics of the food chain are depicted by both the Hastings–Powell model and the chemostat model, with synchrony measured by the Pearson correlation coefficient. We use the density-independent dispersal in the system as a baseline for comparison. Results show that the density-independent dispersal of a species in the system can promote its dynamic synchrony. Dispersal of intermediate species in the tri-trophic food chain is the strongest synchronizer. In contrast, the density-dependent PP and PE of intermediate species can desynchronize the system. Highly synchronized dynamics emerged when the basal species has a strong PE strategy or when the top species has a moderate PP strategy. Our results reveal the complex relationship between density-dependent dispersal and spatial synchrony in tri-trophic systems.     

Primary Seed Dispersal by Red Howler Monkeys and the Effect of Defecation Patterns on the Fate of Dispersed Seeds1

The effectiveness of a seed disperser depends on the quantity and quality of dispersal. The quality of dispersal depends in large part on factors that affect the post–dispersal fate of seeds, and yet this aspect of dispersal quality is rarely assessed. In the particular case of seed dispersal through endozoochory, the defecation pattern produced has the potential of affecting the fate of dispersed seeds and consequently, dispersal quality and effectiveness. In this study, I assessed the effects of dung presence and dung/seed densities on seed predation by rodents and secondary dispersal by dung beetles. In particular, I compared seed fates in clumped defecation patterns, as those produced by howler monkeys, with seed fates in scattered defecation patterns, as those produced by other frugivores. I also determined the prevalence of red howler monkeys (Alouatta seniculus) as seed dispersers at the plant community level in Central Amazonia by determining the number of species they dispersed in a 25–month period. I found that dung presence and amount affected rodent and dung beetle behavior. Seed predation rates were higher when dung was present, and when it was in higher densities. The same number of seeds was buried by dung beedes, in dumped versus scattered defecation patterns, but more seeds were buried when they were inside large dung–piles versus small piles. Seed density had no effect on rodent or dung beetle behavior. Results indicate that caution should be taken when categorizing an animal as a high or low quality seed disperser before carefully examining the factors that affect the fate of dispersed seeds. Red howler monkeys dispersed the seeds of 137 species during the study period, which is the highest yet reported number for an Alouatta species, and should thus be considered highly prevalent seed dispersers at the plant community level in Central Amazonian terra firme rain forests.     

Density dependence and habitat preference shape seedling survival in a subtropical forest in central China

Aims Seedlings are vulnerable to many kinds of fatal abiotic and biotic agents, and examining the causes of seedling dynamics can help understand mechanisms of species coexistence. To disentangle the relative importance of neighborhood densities, habitat factors and phylogenetic relatedness on focal seedling survival, we monitored the survival of 5306 seedlings of 104 species>15 months. We address the following questions: (i) How do neighborhood densities, habitat variables and phylogenetic relatedness affect seedling survival? What is the relative importance of conspecific densities, habitat variables and phylogenetic relatedness to seedling survival? (ii) Does the importance of the neighborhood densities, habitat variables and phylogenetic relatedness vary among growth forms, leaf habits or dispersal modes? Specially, does the conspecific negative density dependence inhibit tree and deciduous seedlings more compared with shrub and evergreen species? Does density dependence affect the wind and animal-dispersed species equally?Methods We established 135 census stations to monitor seedling dynamics in a 25-ha subtropical forest plot in central China. Conspecific and heterospecific seedling density in the 1-m 2 seedling plot and adult basal area within a 20-m radius provided neighborhood density variables. Mean elevation, convexity and aspect of every 5- × 5-m grid with seedling plots were used to quantify habitat characteristics. We calculated the relative average phylodiversity between focal seedling and heterospecific neighbors to quantify the species relatedness in the neighborhood. Eight candidate generalized linear mixed models with binominal error distribution were used to compare the relative importance of these variables to seedling survival. Akaike's information criteria were used to identify the most parsimonious models.Important findings At the community level, both the neighborhood densities and phylogenetic relatedness were important to seedling survival. We found negative effects of increasing conspecific seedlings, which suggested the existence of species-specific density-dependent mortality. Phylodiversity of heterospecific neighbors was negatively related to survival of focal seedlings, indicating similar habitat preference shared among phylogenetically closely related species may drive seedling survival. The relative importance of neighborhood densities, habitat variables and phylogenetic relatedness varied among ecological guilds. Conspecific densities had significant negative effect for deciduous and wind-dispersed species, and marginally significant for tree seedlings>10cm tall and animal-dispersed species. Habitat variables had limited effects on seedling survival, and only elevation was related to the survival of evergreen species in the best-fit model. We conclude that both negative density-dependent mortality and habitat preference reflected by the phylogenetic relatedness shape the species coexistence at seedling stage in this forest.     

Variation within and between Closely Related Species Uncovers High Intra-Specific Variability in Dispersal

Mounting evidence shows that contrasting selection pressures generate variability in dispersal patterns among individuals or populations of the same species, with potential impacts on both species dynamics and evolution. However, this variability is hardly considered in empirical works, where a single dispersal function is considered to adequately reflect the species-specific dispersal ability, suggesting thereby that within-species variation is negligible as regard to inter-specific differences in dispersal abilities. We propose here an original method to make the comparison of intra- and inter-specific variability in dispersal, by decomposing the diversity of that trait along a phylogeny of closely related species. We used as test group European butterflies that are classic study organisms in spatial ecology. We apply the analysis separately to eight metrics that reflect the dispersal propensity, the dispersal ability or the dispersal efficiency of populations and species. At the inter-specific level, only the dispersal ability showed the signature of a phylogenetic signal while neither the dispersal propensity nor the dispersal efficiency did. At the within-species level, the partitioning of dispersal diversity showed that dispersal was variable or highly variable among populations: intra-specific variability represented from 11% to 133% of inter-specific variability in dispersal metrics. This finding shows that dispersal variation is far from negligible in the wild. Understanding the processes behind this high within-species variation should allow us to properly account for dispersal in demographic models. Accordingly, to encompass the within species variability in life histories the use of more than one value per trait per species should be encouraged in the construction of databases aiming at being sources for modelling purposes.     

Density-dependent dispersal and relative dispersal affect the stability of predator-prey metacommunities

Although density-dependent dispersal and relative dispersal (the difference in dispersal rates between species) have been documented in natural systems, their effects on the stability of metacommunities are poorly understood. Here we investigate the effects of intra- and interspecific density-dependent dispersal on the regional stability in a predator-prey metacommunity model. We show that, when the dynamics of the populations reach equilibrium, the stability of the metacommunity is not affected by density-dependent dispersal. However, the regional stability, measured as the regional variability or the persistence, can be modified by density-dependent dispersal when local populations fluctuate over time. Moreover these effects depend on the relative dispersal of the predator and the prey. Regional stability is modified through changes in spatial synchrony. Interspecific density-dependent dispersal always desynchronizses local dynamics, whereas intraspecific density-dependent dispersal may either synchronize or desynchronize it depending on dispersal rates. Moreover, intra- and interspecific density-dependent dispersal strengthen the top-down control of the prey by the predator at intermediate dispersal rates. As a consequence the regional stability of the metacommunity is increased at intermediate dispersal rates. Our results show that density-dependent dispersal and relative dispersal of species are keys to understanding the response of ecosystems to fragmentation.     

A multivariate analysis of fine-scale species density in the plant communities of a saltwater lagoon – the importance of disturbance intensity

Several factors might influence an organism's tendency or willingness to leave a given patch. One such factor is conspecific density, which may affect the per capita emigration rate. Some previous field studies on butterflies have reported positively density-dependent dispersal (emigration increases with population density) whereas the opposite, negatively density-dependent dispersal, has been found in other species. We investigated the effect of conspecific density on both the tendency to cross a patch boundary and within-patch mobility in Melitaea cinxia , by experimentally manipulating density in large outdoor cages divided into two habitat patches, separated by a barrier of unsuitable habitat. In contrast to previous results for M. cinxia , we found that the butterflies moved away from a patch at higher rates in high conspecific density (positively density-dependent emigration). The within-patch mobility, measured as the distance travelled per time unit, was however unaffected by butterfly density. A possible explanation for the seeming discrepancy with previous results could be that we used higher butterfly densities. For species with fluctuating population dynamics, such as M. cinxia , dispersal activity both at low and at high local density will be important for population phenomena such as fluctuations in distributional range over good and bad years.     

Analysis of the specificity of three root-feeders towards grasses in coastal dunes

Among the root-feeding nematodes that accumulate in the rhizosphere of grasses in European dunes, the genus Pratylenchus is of special relevance given its diversity and distribution. Although different species of Pratylenchus have been reported in dune grasses, the specificity towards dune plants, a fundamental aspect of the biology of the species, has hitherto not been studied. Two inoculation experiments using different combinations of grasses and nematodes were performed. The multiplication and the effect on plant growth of P. dunensis and P. brzeskii, two species which only occur in dune areas was compared with that of P. penetrans, a broad host-range species. The three Pratylenchus spp. could multiply under all hosts; however, there was a clear host-dependent response. The species-specific response observed might account for the shift of Pratylenchus spp. detected in the field. Although, a negative effect on the growth of A. arenaria was demonstrated for the three nematode species, different densities were needed to observe the same effects in plant biomass which point at nematode-specific tolerance. While the typical dune species needed very high densities to produce damage, P. penetrans needed very few specimens. The results obtained indicate that species with similar feeding adaptations show very different multiplication abilities on co-occurring hosts, an aspect that is usually overlooked for belowground herbivores in natural systems. The obtained results might suggest a coevolutionary relationship between specific nematode species and Ammophila arenaria.     

Effects of body size and resource availability on dispersal in a native and a non-native estuarine snail

I manipulated snail densities of two coexisting species of salt marsh snail, Cerithidea californica Haldeman (native) and Batillaria attramentaria Sowerby (non-indigenous) to investigate how resource levels set by intraspecific competition may influence dispersal rates. I used two distinct size classes of the snails (mature and immature) to determine if the effects of competition on dispersal differed between developmental stages of a consumer. Dispersal attempts were measured within enclosure pens by counting snails climbing the sides of the enclosure. The influence of snail density per se and resource levels (which were set by snail densities) on dispersal rates were separated by comparing responses of snails to density before and after resources became depleted. For large snails of both species, dispersal increased as resource levels decreased, supporting the hypothesis that competition influences dispersal rates. Small snails of both species, in contrast, always dispersed at relatively higher rates than larger individuals, but were not influenced by variation in resource levels. This result corroborates other studies that have shown reduced competition in these species at smaller size, and suggests that another mechanism, such as genetically coded behavior to disperse when young, influences their behavior. Previous experiments demonstrated Batillaria's superior resource conversion efficiency; therefore, I had hypothesized that for any given resource level, Cerithidea would disperse more, because it was more affected by resource availability. Adult Batillaria, however, responded more sensitively to resource levels (i.e., dispersed more at any given resource level) than Cerithidea. This counter-intuitive result illustrates the potential importance of genetic limitations on behavioral responses available to a species. Constraints on behavioral responses may have been accentuated since Batillaria is a non-indigenous species whose evolved behavioral responses are not necessarily adapted to its present, non-native environment.     

Differential effects of food availability on population growth and fitness of three species of estuarine, bacterial-feeding nematodes

The significance of bottom-up controls on biological communities has been a long-standing topic of interest in ecology. However, before environmental effects on communities can be properly assessed, a thorough knowledge of the individual species' responses is required. We studied effects of food availability on population development and on different life-history traits in three species of bacterial-feeding nematodes, Diplolaimelloides oschei, Diplolaimelloides meyli (both Monhysteridae) and Pellioditis marina (Rhabditidae), which co-occur on macrophyte detritus in the Westerschelde Estuary (SW Netherlands). The bacteria Escherichia coli was offered in five food-availability treatments corresponding to initial cell densities from 3 × 10¹⁰ cells ml^− 1 to 3 × 10⁷ cells ml^− 1. The three bacterial-feeding nematode species studied here showed differential responses to food availability, which agreed with the general idea that Rhabditidae have extreme colonization abilities under very high food availability, while Monhysteridae tend to have a somewhat slower population development and comparatively lower food requirements. Several life-history traits, including juvenile mortality and development time, did not exhibit a clear food-availability dependence, but bioenergetics-related parameters did. Results on the F1 generation may, however, be affected by strong maternal effects on life-history traits of their progeny. Patterns of food-availability dependence of population increase and size at maturity were similar in P. marina. Both Diplolaimelloides species, however, exhibited a large body size at maturity but a very low population increase at the highest food availability, suggesting a trade-off between biomass and reproduction. Comparison with published data on other nematode species reveals that nematode responses to food availability as well as to other environmental factors are highly species-specific.