Influence of seston quantity and quality on growth of tropical cladocerans.

The seston of the small, shallow, and tropical lake Monte Alegre was tested for quantity and quality for cladocerans by growth bioassays, which were carried out in spring (Daphnia gessneri and Moina micrura), summer (D. gessneri, M. micrura, Ceriodaphnia cornuta, and Simocephalus mixtus), and winter (D. gessneri and D. ambigua). Cohorts of newborns originating from ovigerous females collected in the lake or from laboratory cultures were submitted, at a room temperature of 23 degrees C to the following treatments: (1) the chlorophytes Ankistrodesmus falcatus and/or Scenedesmus spinosus; (2) lake seston; and (3) lake seston + chlorophytes. Growth rate, clutch size, and fecundity were evaluated. Seston alone was not the best food for promoting cladoceran growth. There were seasonal differences in food quantity and quality with spring and summer seston being better for growth than that of the winter. Adding chlorophytes to the seston increased clutch size and fecundity for most species in summer and winter, but not in spring. Energy limitation seems to be the most important factor influencing cladoceran growth in summer and especially in winter.     

The susceptiblity of Keratella cochlearis to interference from small cladocerans

SUMMARY. 1. Even at high population densities (300–2000 ind. 1⁻¹), only one of five small cladoceran species (adult body length <1 mm) significantly suppressed population growth of the rotifer Keratella cochlearis through interference (encounter) competition. At 500 ind. H, adults of D. ambigua (0.96 mm body length) imposed an instantaneous per capita death rate of 0.21 day⁻¹ on this rotifer. These short-term experiments may have underestimated cladoceran interference because newborn rotifers were rarely present.
2. Newborn rotifers (<12 h old) were much more susceptible than adult rotifers (> 24 h old) to interference from Ceriodaphnia dubia. All of the small cladoceran species tested were very much less likely than large Daphnia (body lengths >1.2 mm) to interfere with K. cochlearis , but perhaps at high population densities they could suppress population growth of susceptible rotifer species by damaging, and possibly eating, relatively small and soft-bodied newborn individuals.
3. K. cochlearis of the tecta form, without a posterior spine, produced offspring of the typica form, with a posterior spine, in the presence of C. dubia. This developmental response is stimulated by at least several, and possibly all, cladocerans and probably reduces the susceptibility of the rotifer to cladoceran interference.     

Enemy-mediated apparent competition: empirical patterns and the evidence

Apparent competition arises when two victim species negatively affect each other (−,  −) by enhancing the equilibrium density or changing the foraging behaviour of a shared natural enemy. Shared enemies can also mediate non-reciprocal (−,  0) indirect effects, i.e. indirect amensalism, whenever one prey species is not affected by the presence of alternative prey. We review 34 studies on terrestrial and freshwater systems to evaluate the extent to which apparent competition has been perceived as a reciprocal (−,  −) or non-reciprocal (−,  0) interaction. We found only three studies showing reciprocal effects between apparent competitors. Indirect amensalism was documented in 10 studies and could be inferred for 16 other cases (76% in total). The remaining five studies provided insufficient data to determine the form of indirect interaction. The apparent prevalence of non-reciprocal enemy-mediated interactions resembles that observed for resource-based interspecific competition. Amensal indirect effects via shared predation may result from differences in population size, nutritional value, susceptibility to attack, or asynchronous dynamics of alternative prey, or the predator's feeding preferences. Moreover, experimental protocols may confound the actual form of apparent competition through short-term observations, incomplete designs, or biased consideration of conspicuous interactions, leading to reciprocal effects being overlooked. We conclude that, at present, it is still difficult to determine the relative role of apparent competition vs indirect amensalism in natural food webs because most published studies have failed to document in full interactions via shared enemies.     

An analysis of seasonal dynamics of a mixed population of Daphnia, and the associated cladoceran community

SUMMARY. The dynamics of a mixed population of Daphnia (D. ambigua and D. parvula) are analysed over 9 months in a small eutrophic lake. A spring peak of D. ambigua occurred shortly after ice break-up. Decline of D. ambigua coincided with a high occurrence of sexual reproduction, the appearance of D. parvula which did not become abundant, and a substantial increase in Ceriodaphnia quadrangula . A third cladoceran, Bosmina longirostris , was abundant before and after the period of numerical dominance by D. ambigua , but declined to very low numbers coincident with the increase in D. ambigua.
Estimates of egg hatching rate at various temperatures demonstrate that these three cladocerans have different rate responses to temperature, and that seasonal succession of species is predictable from temperature alone. The importance of temperature versus species interactions in controlling species succession is discussed.     

Predator-Mediated Indirect Effects of Snowshoe Hares on Dall's Sheep in Alaska

ABSTRACT Indirect interactions among species can strongly influence population dynamics and community structure but are often overlooked in management of large mammals. We estimated survival of Dall's sheep (Ovis dalli) in the central Alaska Range, USA, during years of differing snowshoe hare (Lepus americanus) abundance to test whether indirect interactions with a cyclic hare population affect Dall's sheep either negatively, by subsidizing predators (apparent competition), or positively, by diverting predation (apparent commensalism). Annual survival of adult female sheep was consistently high (0.85 for all yr and age classes combined). In contrast, annual estimates of lamb survival ranged from 0.15 to 0.63. The main predators of lambs were coyotes (Canis latrans) and golden eagles (Aquila chrysaetos), which rely on hares as their primary food and prey on lambs secondarily. Coyotes and eagles killed 78% of 65 radiocollared lambs for which cause of death was known. Lamb survival was negatively related to hare abundance during the previous year, and lamb survival rates more than doubled when hare abundance declined, supporting the hypothesis of predator-mediated apparent competition between hares and sheep. However, stage-specific predation and delays in predator responses to changes in hare numbers led to a positive relationship between abundance of adult Dall's sheep and hares. Lacking reliable estimates of survival, a manager might erroneously conclude that hares benefit sheep. Thus, support for different indirect effects can be obtained from different types of data, which demonstrates the need to determine the mechanisms that create indirect interactions. Long-term survey data suggest that predation by coyotes is limiting this sheep population below levels typical when coyotes were rare or absent. Understanding the nature of indirect interactions is necessary to effectively manage complex predator–prey communities.     

Dynamics of the exotic Daphnia lumholtzii and native macro-zooplankton in a subtropical chain-of-lakes in Florida, U.S.A.

1. Seasonal dynamics of the exotic Daphnia lumholtzii and native macro-zooplankton species were studied for 2 years in six inter-connected lakes in Florida, U.S.A. The lakes ranged widely in pH, colour and trophic status, and were dominated by copepods.
2. All six lakes contained both D. lumholtzii and the native D. ambigua , but the two species did not overlap in time. D. ambigua was dominant in autumn–spring, coinciding with lower water temperature, higher transparency and lower nutrient and chlorophyll a (Chl a ) concentrations than in summer, when D. lumholtzii was dominant.
3. Based on the field observations, temperature optima were 24 °C for D. ambigua and 29 °C for D. lumholtzii , suggesting that temperature plays a role in determining dominance among the daphnids of subtropical Florida lakes.
4.  D. lumholtzii has not displaced native cladocerans but occupies a 'vacant' seasonal niche, unexploited due to the inability of native taxa to tolerate high temperature. Furthermore, D. lumholtzii did not significantly alter algal–zooplankton interactions. There was evidence of top–down control by grazing, but it was primarily attributable to the native taxa.     

Genotype‐by‐environment interaction and the Dobzhansky–Muller model of postzygotic isolation

The Dobzhansky–Muller (D–M) model of reproductive isolation (RI) posits that hybrid sterility and inviability result from negative epistatic interactions between alleles at a minimum of two genes. This standard model makes several implicit assumptions, including a lack of environmental effects and genotype‐by‐environment interactions (GEI) involving hybrid sterility and hybrid inviability loci. Here we relax this assumption of the standard D–M model. By doing so, several patterns of the genetic architecture of RI change. First, a novel single‐locus model of postzygotic RI emerges. Several indirect lines of evidence are discussed in support of the model, but we conclude that this new single‐locus model is currently no more supported than previous ones. Second, when multilocus D–M models incorporating GEI are considered, we find that the number of potential negative epistatic interactions increases dramatically over the number predicted by the standard D–M model, even when only the most simple case of two‐allele interactions are considered. Third, these multilocus models suggest that some previous generalizations about the evolutionary genetics of postzygotic RI may not necessarily hold. Our findings also suggest that the evolution of postzygotic RI may be more likely when the expression of traits driving speciation is affected by the environment, since there appears to be a greater spectrum of potential hybrid incompatibilities under the D–M model incorporating GEI.     

Predator-mediated interactions among the seeds of desert plants

In theory, seed predators are capable of inducing indirect interactions among the seeds they consume. However, empirical evidence of predator-mediated interactions among seeds is rare. Rodents in the Heteromyidae are highly granivorous and therefore likely to induce indirect interactions among the seeds of desert plants. The indirect interactions may be in the form of apparent competition and apparent mutualism between seeds within a patch. Apparent competition exists when the survival of seeds of a focal species is lessened because of the presence of additional seeds of other species in the patch. Apparent mutualism exists when the presence of the other seeds results in an increase in survival of seeds of the focal species. By measuring seed removal from trays placed in the field, apparent competition between the seeds of several plant species was detected. Apparent mutualism might also exist, but there was no strong evidence of it. Apparent competition appeared most likely to occur among the species whose seeds were the most heavily predated. For instance, predation on seeds of Astragalus cicer, Oryzopsis hymenoides, and Sphaeralcea coccinea was substantial with more than 50% of the seeds removed from the trays, on average. The intensity of apparent competition (measured by the indirect effect, IS) between these species and two others was significant; IS ranged from –0.02 to –0.39 on a scale of 0 to –1. This indicates that, in some communities, indirect effects are most likely to exist when direct effects are strong. Received: 5 August 1999 / Accepted: 2 March 2000     

不同基质中水鳖对共存的穗花狐尾藻和金鱼藻的影响

研究设置了营养水平不同的3种基质,在有或无浮水植物水鳖共存的情况下,混合种植营养吸收方式不同的两种沉水植物（仅从水中吸收营养的金鱼藻和从水和基质都能吸收营养的穗花狐尾藻）,探讨在不同的基质中浮水植物对沉水植物的影响。结果显示:实验中水鳖的存在均未对共存的两种沉水植物的生长产生抑制作用。当水鳖覆盖水面时,与无根的金鱼藻竞争光照和水体营养,促使其增加分枝数和茎长,并增加叶生物量的分配,最终导致水鳖的共存促进了金鱼藻的生物量积累。但与金鱼藻不同,有根的穗花狐尾藻可通过基质吸收营养,与共存的水鳖之间只存在光照竞争,使穗花狐尾藻仅以改变植株形态来适应浮水植物的表层覆盖,生物量积累并未发生显著变化。与沙处理相比,水鳖在营养水平较高的泥处理和泥沙混合处理中生长更旺盛,基质没有显著影响沉水植物的生物量和分株数。因此,在退化湿地生态系统中恢复水生植物时,为了提高群落的物种多样性,可以将沉水植物与少量的浮水植物组合配置,少量的浮水植物不会对沉水植物的生长带来负面影响。     

Stability of a diamond-shaped module with multiple interaction types

Indirect interactions among species emerge from the complexity of ecological networks and can strongly affect the response of communities to disturbances. To determine these indirect interactions and understand better community dynamics, ecologists focused on the interactions within small sets of species or modules. Thanks to their analytical tractability, modules bring insights on the mechanisms occurring in complex interaction networks. So far, most studies have considered modules with a single type of interaction although numerous species are involved in mutualistic and antagonistic interactions simultaneously. In this study, we analyse the dynamics of a diamond-shaped module with multiple interaction types: two resource species sharing a mutualist and a consumer. We describe the different types of indirect interaction occurring between the resource species and the conditions for a stable coexistence of all species. We show that the nature of indirect interactions between resource species (i.e. apparent facilitation, competition or antagonism), as well as stable coexistence, depend on the species generalism and asymmetry of interactions, or in other words, on the distribution of interaction strengths among species. We further unveil that a balance between mutualistic and antagonistic interactions at the level of resource species favours stable coexistence, and that species are more likely to coexist stably if there is apparent facilitation between the two resource species rather than apparent competition. Our results echo existing knowledge on the trophic diamond-shaped module, and confirm that our understanding of communities combining different interaction types can gain from module analyses.     

Associational resistance or susceptibility: the indirect interaction between chemically‐defended and non‐defended herbivore prey via a shared predator

Many organisms possess chemical defences against their natural enemies, which render them unpalatable or toxic when attacked or consumed. These chemically‐defended organisms commonly occur in communities with non‐ or less‐defended prey, leading to indirect interactions between prey species, mediated by natural enemies. Although the importance of enemy‐mediated indirect interactions have been well documented (e.g. apparent competition), how the presence of prey chemical defences may affect predation of non‐defended prey in terrestrial communities remains unclear. Here, an experimental approach was used to study the predator‐mediated indirect interaction between a chemically‐defended and non‐defended pest aphid species. Using laboratory‐based mesocosms, aphid community composition was manipulated to include chemically‐defended (CD) aphids Brevicoryne brassicae, non‐defended (ND) aphids Myzus persicae or a mixed assemblage of both species, on Brassica oleracea cabbage plants, in the presence or absence of a shared predator (Chrysoperla carnea larvae). Aphid population growth rates, aphid distributions on host plants and predator growth rates were measured. In single‐species treatments, C. carnea reduced M. persicae population growth rate, but had no significant impact on B. brassicae population growth rate, suggesting B. brassicae chemical defences are effective against C. carnea. Chrysoperla carnea had no significant impact on either aphid species population growth rate in mixed‐species treatments. Myzus persicae (ND) therefore experienced reduced predation in the presence of B. brassicae (CD) through a predator‐mediated indirect effect. Moreover, predator growth rates were significantly higher in the M. persicae‐only treatments than in either the B. brassicae‐only or mixed‐species treatments, suggesting predation was impaired in the presence of B. brassicae (CD). A trait‐mediated indirect interaction is proposed, consistent with associational resistance, in which the predator, upon incidental consumption of chemically‐defended aphids is deterred from feeding, releasing non‐defended aphids from predatory control.     

Direct and indirect effects in microcosm communities of protists

Increased complexity in biological communities can increase the variety of interactions among species, but the relative strengths and long-term consequences of various direct and indirect interactions require further investigation. I studied interactions among four species of protists by monitoring their population dynamics when they were cultured either together or in seven different subset communities. Two protists were bacterivores (Chilomonas and Tetrahymena) and two were predators (Actinosphaerium and Euplotes). Actinosphaerium was omnivorous, and could eat both predatory Euplotes and bacterivores. Three indirect effects occurred among the four species of protists, including indirect facilitation of one predator by the other, apparent competition between bacterivores, and indirect facilitation of one bacterivore by the omnivorous predator. Community structure and invasibility depended on both direct and indirect effects; thus both can be mechanisms for assembly rules.     

Cladocerans (Branchiopoda) of a tropical estuary in Brazil.

The study was conducted in the Capibaribe river estuary in Recife (Brazil) to assess the role played by cladocerans in a eutrophic environment. Samplings were carried out monthly at 4 fixed stations along the estuary from July 1987 to June 1988. Collections were made with a plankton net of 65 micrometers mesh size. Six cladocerans species were registered: Penilia avirostris, Diaphanosoma spinulosum, Chlydorus barroisi, Ceriodaphnia rigaudi, Ilyocryptus spinifer, and Moina micrura. The most frequent species was Moina micrura with 49%. The only true marine species was Penilia avirostris, which was registered at station 1 (close to the river mouth), during high and low tide and in the dry season. The Cladocera average density decreased from 329 ind.m'3 (August 1987, high tide) to 2 ind.m(-3) (March 1988, high tide) due to the high load of organic and chemical pollution received by the estuary. At all stations, D. spinulosum, M. micrura, and C. barroisi occurred with a wide distribution, mainly during the rainy season. C. rigaudi and I. spinifer were rare, occurring only during the rainy season. Cladocerans played an important role in the food webs of the plankton community of the Capibaribe tropical estuary and the dominance of a few small species indicated a hypereutrophic environment. A high level of disturbance was indicated by the decline in diversity of specialized species and the increase in abundance of opportunistic species like M. micrura.     

The effect of the Holling type II functional response on apparent competition

This article analyzes the classical 2-resource-1-consumer apparent competition community module with the Holling type II functional response. Two types of resource regulation (top-down vs. combined top-down and bottom-up) and two types of consumer behaviors (inflexible consumers with fixed preferences for resources vs. adaptive consumers) are considered. When resources grow exponentially and consumers are inflexible foragers, one resource is always outcompeted due to strong apparent competition. Density dependent resource growth relaxes apparent competition so that resources can coexist. As multiple attractors (either equilibria or limit cycles) coexist, population dynamics and community composition depend on initial population densities. Population dynamics change dramatically when consumers forage adaptively. In this case, the results both for top-down, and combined top-down and bottom-up regulation are similar and they show that species persistence occurs for a much larger set of parameter values when compared with inflexible consumers. Moreover, population dynamics will be chaotic when resource carrying capacities are high enough. This shows that adaptive consumer switching can destabilize population dynamics.     

Trophic and non-trophic pathways mediate apparent competition through post-dispersal seed predation in a Patagonian mixed forest

Post‐dispersal predation can be a major source of seed loss in temperate forests. Little is known, however, about how predator‐mediated indirect interactions such as apparent competition alter survival patterns of canopy tree seeds. Understorey plants may enhance tree seed predation by providing sheltered habitat to granivores (non‐trophic pathway). In addition, occurrence of different tree seeds in mixed patches may lead to short‐term apparent competition between seed types, because of the granivores’ foraging response to changes in food patch quality (trophic pathway). We hypothesised that understorey bamboo cover and mixing of seed species in food patches would both increase tree seed predation in a Nothofagus dombeyi?Austrocedrus chilensis forest in northern Patagonia, Argentina. Seed removal experiments were conducted for three consecutive years (2000–2002) differing in overall granivory rates. Seed patch encounter and seed removal rates were consistently higher for the larger and more nutritious Austrocedrus seeds than for the smaller Nothofagus seeds. Seed removal was greater beneath bamboo than in open areas. This apparent competition pathway was stronger in a low‐predation year (2000) than in high‐predation years (2001–2002), suggesting a shift in microhabitat use by rodents. Patch composition had a significant, though weaker, impact on seed survival across study years, whereas seed density per patch enhanced encounter rates but did not influence seed removal. Removal of the less‐preferred Nothofagus seeds increased in the presence of Austrocedrus seeds, but the reciprocal indirect effect was not observed. However, this non‐reciprocal apparent competition between seed species was only significant in the high‐predation years. Our study shows that granivore‐mediated indirect effects can arise through different interaction pathways, affecting seed survival patterns according to the predator's preference for alternative seed types. Moreover, results indicate that the occurrence and relative strength of trophic vs non‐trophic pathways of apparent competition may change under contrasting predation scenarios.     

Effects of the blue-green alga Microcystis aeruginosa on zooplankton competitive relations

Summary Field distribution patterns and laboratory feeding experiments have suggested that blooms of colonial blue-green algae strongly inhibit relatively large-bodied daphnid cladocerans. We conducted laboratory experiments to test the hypothesis that blooms of the colonial blue-green alga Microcystis aeruginosa would shift competitive dominance away from large-bodied daphnid cladocerans toward smaller-bodied cladocerans, copepods, and rotifers. In laboratory competition experiments, increasing the proportion of M. aeruginosa in the algal food supply resulted in a shift from dominance by the relatively largebodied cladoceran Daphnia ambigua to dominace by the copepod Diaptomus reighardi. The small-bodied cladoceran Bosmina longirostris was always numerically heavily dominant over D. ambigua, but its estimated population biomasses were only slightly higher than those of D. ambigua. Daphnia ambigua consistently outcompeted the rotifer Brachionus calyciflorus. Our results demonstrate that blooms of M. aeruginosa can alter zooplankton competitive relations in laboratory experiments, favoring small-bodied cladocerans and copepods at the expense of large-bodied cladocerans. However, contrary to predictions, blooms of M. aeruginosa did not improve the competitive ability of rotifers.     

Biotic indirect effects: a neglected concept in invasion biology

Indirect effects involve more than two species and are defined as how one species alters the effect that another species has on a third. These complex interactions are often overlooked in studies of interactions between alien and native species, and their role in influencing biological invasions has been rarely considered. Based on a comprehensive review of the invasion biology literature, we examine the evidence for the occurrence of four of the most commonly documented indirect effects (apparent competition, indirect mutualism/commensalism, exploitative competition, and trophic cascades) in the invasion process. Studies investigating indirect effects in the context of invasion biology are relatively rare, but have been increasing in recent years, and there are sufficient examples to indicate that this kind of interaction is likely to be more common than is currently recognized. Whether indirect interactions are mediated by an alien or a native species, and whether they occur between ecologically similar or dissimilar alien and native species, depends in part on the type of interaction considered and no predictable patterns were detected in the literature. Further empirical studies will help to elucidate such patterns. At this stage, the inherent unpredictability of indirect interactions means that their impacts in relation to invasions are particularly challenging for land managers to deal with, and their role in invasions is a complex, but is a valuable area of investigation for researchers.     

Interspecific synchrony of seed rain shapes rodent‐mediated indirect seed–seed interactions of sympatric tree species in a subtropical forest

Animal‐mediated indirect interactions play a significant role in maintaining the biodiversity of plant communities. Less known is whether interspecific synchrony of seed rain can alter the indirect interactions of sympatric tree species. We assessed the seed dispersal success by tracking the fates of 21 600 tagged seeds from six paired sympatric tree species in both monospecific and mixed plots across 4 successive years in a subtropical forest. We found that apparent mutualism was associated with the interspecific synchrony of seed rain both seasonally and yearly, whereas apparent competition or apparent predation was associated with interspecific asynchrony of seed rain either seasonally or yearly. We did not find consistent associations of indirect interactions with seed traits. Our study suggests that the interspecific synchrony of seed rain plays a key role in the formation of animal‐mediated indirect interactions, which, in turn, may alter the seasonal or yearly seed rain schedules of sympatric tree species.     

铜绿微囊藻对萼花臂尾轮虫与大型溞竞争关系的影响

为探究铜绿微囊藻(Microcystis aeruginosa)浓度变化对浮游动物竞争关系的影响,通过控制实验法,评估了在3个铜绿微囊藻浓度梯度下,萼花臂尾轮虫(Brachionus calyciflorus)和大型溞(Daphnia magna)之间的种间竞争关系。结果表明不同浓度铜绿微囊藻对萼花臂尾轮虫、大型溞的增长及二者种间竞争影响具有差异,并且在3种铜绿微囊藻浓度下均以大型溞为主要优势类群。低浓度(5×10⁴ cells/m L)铜绿微囊藻仅促进大型溞种群增长(P<0.01),大型溞占据主要优势地位;中浓度(1×10⁵ cells/m L)铜绿微囊藻对萼花臂尾轮虫和大型溞增长均有显著影响(P<0.01),在此浓度下大型溞在种群竞争中依旧占优势地位,使得萼花臂尾轮虫种群衰亡;在高浓度铜绿微囊藻(5×10⁵ cells/m L)环境中种群生长均受到抑制(P<0.01),在共培养体系中仅大型溞种群存活。在无其他外在影响因素存在时,实验结果显示在不同浓度的铜绿微囊藻下,大型溞均占优势,说明铜绿微囊藻的浓...     

A shift from exploitation to interference competition with increasing density affects population and community dynamics

Intraspecific competition influences population and community dynamics and occurs via two mechanisms. Exploitative competition is an indirect effect that occurs through use of a shared resource and depends on resource availability. Interference competition occurs by obstructing access to a resource and may not depend on resource availability. Our study tested whether the strength of interference competition changes with protozoa population density. We grew experimental microcosms of protozoa and bacteria under different combinations of protozoan density and basal resource availability. We then solved a dynamic predator–prey model for parameters of the functional response using population growth rates measured in our experiment. As population density increased, competition shifted from exploitation to interference, and competition was less dependent on resource levels. Surprisingly, the effect of resources was weakest when competition was the most intense. We found that at low population densities, competition was largely exploitative and resource availability had a large effect on population growth rates, but the effect of resources was much weaker at high densities. This shift in competitive mechanism could have implications for interspecific competition, trophic interactions, community diversity, and natural selection. We also tested whether this shift in the mechanism of competition with protozoa density affected the structure of the bacterial prey community. We found that both resources and protozoa density affected the structure of the bacterial prey community, suggesting that competitive mechanism may also affect trophic interactions.