Effects of hydroperiod and predation on a Mississippi River floodplain invertebrate community

 The objective of this study was to determine if pond permanence and vertebrate predation (by fish and waterfowl) affect invertebrate community structure in the mudflat habitat of floodplain ponds. Invertebrate communities were studied for 1 year in four Mississippi River floodplain ponds with different hydroperiods. Pond 1 experienced five dry periods, pond 2 experienced four, pond 3 dried once, and standing water remained in pond 4 for the entire year. Vertebrate predator exclusion treatments (all access, no access, small-fish access and cage controls) were placed in all ponds. As pond duration increased, predatory invertebrate richness and abundance increased while overall invertebrate richness and abundance decreased. With the exception of the cladoceran Diaphanosoma, all commonly encountered taxa were strongly affected by pond permanence in terms of abundance, biomass and, generally, individual biomass. Taxa were nearly early divided between those that were more abundant in less permanent ponds and those that were more abundant in longer-duration ponds. Invertebrate taxa richness, abundance, and total biomass were lower in the all-access treatment than in the treatments that restricted predator access, and these effects were stronger in the more permanent ponds. In general, there were no significant differences in responses to the treatments with small-fish access and no access. These results support models that predict relatively weak effects of predation in frequently disturbed habitats. Received: 30 May 1995 / Accepted: 21 June 1996     

Evaluating predation pressure on green treefrog larvae across a habitat gradient

The effect of a predator on the abundance of a prey species depends upon the predators abundance and its ability to capture that prey. The objectives of this research were to evaluate the community structure of predators of green treefrog (Hyla cinerea) tadpoles across habitat types and evaluate the effectiveness of individual predators on H. cinerea tadpoles. Correspondence and cluster analyses of predator frequencies across 23 aquatic habitats indicated that the majority of variance in predator communities was due to a division between permanent and temporary habitats. Experimental work demonstrated that survival of the smallest H. cinerea tadpoles was significantly lower than survival of medium and large tadpoles with the most effective predators, indicating that H. cinerea tadpoles attain a refuge from predation at larger body sizes. We combined the effectiveness of predators in experiments with the abundance of each predator species from the predator community survey to demonstrate that predation pressure on H. cinerea tadpoles is higher in temporary ponds. This pattern may explain in part why this species generally breeds successfully only in permanent habitats. It also confirms that discussions about an increasing gradient of predation pressure from temporary to permanent aquatic habitats should be restricted to individual prey species for which such a gradient has been demonstrated.     

Predator size and phenology shape prey survival in temporary ponds

Theoretical efforts suggest that the relative sizes of predators and their prey can shape community dynamics, the structure of food webs, and the evolution of life histories. However, much of this work has assumed static predator and prey body sizes. The timing of recruitment and the growth patterns of both predator and prey have the potential to modify the strength of predator–prey interactions. In this study, I examined how predator size dynamics in 40 temporary ponds over a 3-year period affected the survival of spotted salamander (Ambystoma maculatum) larvae. Across communities, gape-limited predator richness, but not size, was correlated with habitat duration (pond permanence). Within communities, mean gape-limited predator size diminished as the growing season progressed. This size reduction occurred because prey individuals grew into a body size refuge and because the largest of the predators left ponds by mid-season. Elevated gape-limited predation risk across time and space was predicted by the occurrence of two large predatory salamanders: marbled salamander larvae (Ambystoma opacum) and red-spotted newt adults (Notophthalmus viridescens). The presence of the largest gape-limited predator, A. opacum, predicted A. maculatum larval survival in the field. The distribution of large predatory salamanders among ponds and across time is expected to lead to differing community dynamics and to generate divergent natural selection on early growth and body size in A. maculatum. In general, a dynamic perspective on predator size often will be necessary to understand the ecology and evolution of species interactions. This will be especially true in frequently disturbed or seasonal habitats where phenology and ontogeny interact to determine body size asymmetries. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Population dynamics of Triops cancriformis (Crustacea: Branchiopoda: Notostraca) of the Espolla temporary pond in the northeastern Iberian peninsula

The population dynamics of Triops cancriformis in Espolla temporary pond (NE Iberian peninsula) were studied during 1996 and 1997, which encompassed six flooded periods. Data were collected on each individual's size, sex, and, if female, on number of eggs in the oostegopodes. Male-biased sex ratios were found only in the drying-out phase and variations in fecundity were strongly related to hydroperiod duration. Sex ratio variation during the drying-out phase can be attributed to female mortality because the very low recruitment observed does not support the hypothesis of an increase of males. Two hypotheses are advanced to account for female mortality: (1) differential reproductive effort, and (2) size selective predation by herons. This population is characterised by low values of maximum densities compared with other notostracan populations, and by higher densities in the spring–summer hydroperiods than in the winter ones.     

Sacrificial males: the potential role of copulation and predation in contributing to copepod sex‐skewed ratios

Predation is thought to play a selective role in the emergence of behavioural traits in prey. Differences in behaviour between prey demographics may, therefore, be driven by predation with select components of the population being less vulnerable to predators. While under controlled conditions prey demography has been shown to have consequences for predation success, investigations linking these implications to natural prey population demographics are scarce. Here we assess predator–prey dynamics between notonectid predators (backswimmers) and Lovenula raynerae (Copepoda), key faunal groups in temperate ephemeral pond ecosystems. Using a combination of field and experimental approaches we test for the development and mechanism of predation‐induced sex‐skewed ratios. A natural population of L. raynerae was tracked over time in relation to their predator (notonectid) and prey (Cladocera) numbers. In the laboratory, L. raynerae sex ratios were also assessed over time but in the absence of predation pressure. Predation success and prey performance experiments evaluating differences between L. raynerae male, female, gravid female and copulating pairs exposed to notonectid predation were then examined. Under natural conditions, a female dominated copepod population developed over time and was correlated to predation pressure, while under predator‐free conditions non sex‐skewed prey population demographics persisted. Predator–prey laboratory trials showed no difference in vulnerability and escape performance for male, female and gravid female copepods, but pairs in copula were significantly more vulnerable to predation. This vulnerability was not shared by both sexes, with only female copepods ultimately escaping from successful predation on a mating pair. These results suggest that contact periods during copula may contribute to the development of sex‐skewed copepod ratios over time in ecosystems dominated by hexapod predators. This is discussed within the context of vertebrate and invertebrate predation and how these dissimilar types of predation are likely to have acted as selective pressures for copepod mating systems.     

The influence of predator regime on the behaviour and mortality of a freshwater amphipod, <Emphasis Type="Italic">Gammarus pulex</Emphasis>

In species with restricted dispersal, traits may become genetically fixed leading to local adaptations. Therefore, predator avoidance in a prey species may differ between populations experiencing different predator regimes, but also between sexes within a population due to different vulnerability to predators. In this study we used male and female Gammarus pulex from two different predator regimes: fishless ponds, where invertebrates are the dominant predators and ponds with predatory fish. In the laboratory we examined refuge use, mortality, leaf decomposition rate and pair-formation in G. pulex when exposed to predator cues from either invertebrate predators or fish. Individuals from fish ponds spent more time in refuge and had a higher mortality than those from fishless ponds independent of predator cues. There was no effect of pond predator regime or predator cues on leaf decomposition rates. Further, fewer individuals formed pairs in G. pulex from fish ponds than from fishless ponds. Male G. pulex had a higher mortality and a higher decomposition rate than females independent of predator cues. However, there was no difference in refuge use between sexes. Our study shows that there are general differences in behaviour traits, both between predator regimes and sexes in G. pulex.     

Prey‐mediated changes in the selectivity of the predator Macrolophus pygmaeus (Heteroptera: Miridae)

When foraging in communities with mixed prey, generalist predators may be confronted with prey species that differ in quality, size and mobility and interact with one another. To examine prey selection, predation by Macrolophus pygmaeus (Heteroptera: Miridae) was recorded by providing a diet of either one or two prey species of Myzus persicae (third‐instar nymphs), Aphis gossypii (fourth‐instar nymphs), Trialeurodes vaporariorum (third‐instar nymphs) and Ephestia kuehniella (eggs). In the experiments, prey mobility, prey quality and prey biomass were considered. The biomass consumed by the predator was dependent on the combination of prey species and the quantity of biomass offered. In choice experiments with diets mixed of two prey species at equal densities, the predation to A. gossypii was significantly reduced in the presence of E. kuehniella but the rate of consumption of M. persicae, T. vaporariorum and E.kuehniella was not significantly affected by the coexistence of any other species in the mixed prey diet. When equal amounts of biomass from two prey species were provided in combination, the total consumed biomass was significantly reduced in the mixed prey diets composed of E. kuehniella eggs and aphid nymphs. Thus, under the mixed‐prey situation, prey selection by predators may be affected by interactions among prey species differing in traits such as quality, mobility and size.     

Sequential movement into coastal habitats and high spatial overlap of predator and prey suggest high predation pressure in protected areas

In theory, predators should attempt to match the distribution of their prey, and prey to avoid areas of high predation risk. However, there is a scarcity of empirical knowledge on predator and prey spatial use when both are moving freely in their natural environment. In the current study, we use information collated on a predators’ diet, its population structure, as well as predator and prey relative abundance, and track the movements of predator and prey simultaneously to compare habitat use and evaluate predation pressure. The study was conducted in elasmobranch protected areas of coastal Tasmania, Australia. The species considered were the broadnose sevengill shark Notorynchus cepedianus, the apex predator in the area, and five chondrichthyan prey species. Notorynchus cepedianus and its prey show similar seasonality in the use of these coastal areas: more abundant in warmer months and absent in winter. Predator and prey also showed high spatial overlap and similar habitat use patterns. These similar movement patterns of predator and prey combined with the additional ecological information (diet, population structure of predator, relative abundance of predator and prey) suggests that N. cepedianus move into coastal areas to exploit seasonally abundant prey. Also, while in protected areas, chondrichthyans are subjected to high predation pressure. Overall, results illustrate the value of simultaneously recording and integrating multiple types of information to explore predator–prey relationships and predation pressure.     

Effect of the cyclopoid copepod Mesocyclops thermocyclopoides on the interactions between the predatory rotifer Asplanchna intermedia and its prey Brachionus calyciflorus and B. angularis

In many shallow, eutrophic subtropical ponds, brachionid rotifers are common prey of the predatory copepod Mesocyclops thermocyclopoides. The predatory rotifer Asplanchna intermedia, which is itself a potential prey of the copepod, also feeds preferentially on brachionids. We studied in the laboratory the population dynamics of two mutually competing prey species, Brachionus angularis and B. calyciflorus, in the presence of the two predators A. intermedia and M. thermocyclopoides. The experimental design included separate population dynamics studies with one prey–one predator, two prey–one predator, one prey–two predator, and two prey–two predator systems. These combinations were compared with controls, in which both the prey species (B. angularis and B. calyciflorus) were grown separately and in combination with each other. In the absence of any predator, B. angularis generally eliminated the larger B. calyciflorus. Selective predation by the copepod allowed B. calyciflorus to persist longer in competition with B. angularis. Feeding by M. thermocyclopoides on A. intermedia reduced the predation pressure on B. calyciflorus. However, given enough time, the cyclopoid copepod was able to eliminate both the brachionids as well as the predatory Asplanchna.     

Are behavioural traits in prey sensitive to the risk imposed by predatory fish?

1. Behavioural differences among prey species may result from evolutionary adaptations that facilitate coexistence with different predators and influence vulnerability to predators. It has been hypothesised that prey species modify their behaviour in relation to the risk posed by particular predators. 2. We examined the relationship between anti‐predator behaviour and predation risk in five species of larval odonates in combination with three predatory fish species (perch, gudgeon and rudd) that differ in foraging behaviour. The odonates, Platycnemis pennipes, Coenagrion puella, Lestes sponsa, Sympetrum striolatum and Libellula depressa, differ with regard to their life cycle and habitat, including water depth, occurrence in temporary ponds and co‐existence with fish. 3. The odonate species differed in their response to fish: (i) Two species showed a flexible response. Larval C. puella reduced activity in the presence of fish, regardless of species, whereas L. depressa altered their activity only in the presence of gudgeon. (ii) Independent of fish species, all odonates except L. depressa exhibited spatial avoidance of fish. This was interpreted as a more general anti‐predator response. (iii) In some cases the odonates showed no response to predators and their behaviour was thus independent of predation risk. 4. Our results confirm that all odonates responded to the presence of at least some predatory fish, and that some odonate species discriminated between fish species. However, we found no significant correlation between behavioural modifications and predation risk, indicating that anti‐predator responses and predation risk depend on the particular predator and the species being preyed on.     

Predator‐induced phenotypic plasticity in an arid‐adapted tropical tadpole

Adaptive phenotypic plasticity is widespread and involves diverse phenotypes. Key environmental stressors, such as predation risk, can simultaneously induce changes in multiple traits, but the magnitude of response is dependent upon the environmental conditions. Species that utilize temporary ponds are expected to exhibit stronger predator‐induced responses in the form of morphology than behaviour (i.e. reduced activity) to meet the demands of rapid development by maintaining high foraging activity while reducing predation risk via morphologically plastic traits. In a laboratory experiment, I examined the effects of predator chemical cues and conspecific alarm cues on activity, development and morphology on Leptodactylus bufonius tadpoles. This species has terrestrial oviposition and completes the early part of its development outside of ephemeral and temporary ponds in the Gran Chaco ecoregion of South America. Tadpoles in the predator treatments exhibited both behavioural and morphological predator‐induced plastic responses. Tadpoles tended to possess shorter, deeper tails when exposed to predators. The greatest reduction in activity was observed in tadpoles exposed to both predator and conspecific alarm cues, which subsequently resulted in the slowest development. Temporary and ephemeral pond adapted species with terrestrial oviposition may capitalize on a head start in development by being able to afford reduced growth rates via a reduction in activity. This may occur when the constraints imposed by pond hydroperiod (e.g. risk of pond drying) are relaxed when compared with species with aquatic oviposition, which must undergo all stages of development during the pond's hydroperiod. Thus, in addition to the predator and hydroperiod gradients, examining phenotypically plastic responses along a ‘terrestriality gradient’ in a comparative framework would provide insights as to the costs and benefits of increasing terrestriality in anuran reproductive modes to environmental stressors.     

Body mass relationships affect the age structure of predation across carnivore–ungulate systems: a review and synthesis

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Trait flexibility of generalist invertebrates exposed to contrasting predation and drying stressors

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Effects of predation pressure on the feeding behaviour of the serpa tetra Hyphessobrycon eques (Ostariophysi, Characidae)

The indirect, non-lethal results of predation, such as reduction in feeding time or restraint in seeking sexual partners and/or natural resources, have a drastic effect on prey populations. In this study, we investigated the behaviour of two serpa tetras Hyphessobrycon eques groups, one wild and the other born and raised in captivity, to evaluate how their feeding behaviour is affected by the avian predator, the rufescent tiger-heron Trigrisoma lineatum (using a taxidermy specimen). For a total of 133 observation hours, the feeding behaviour of each fish group was observed according to the absence (control) or presence (treatments) of a predator near the aquarium surface. The results showed that the presence of a predator on the surface inhibits the feeding behaviour of H. eques. The differences observed between the groups are probably due to the fishes experiences with predators. Our results suggest that fish-eating birds may affect prey populations in streams and ponds perhaps more through non-lethal effects, on feeding behaviour for example, than directly through death rates.     

Diet specialization in a fluctuating population of Saduria entomon: a consequence of resource or forager densities?

Intraspecific competition has been shown to favor diet specialization among individuals. However, the question whether the competition takes the form of interference or exploitative in driving diet specialization has never been investigated. We investigated individual diet specialization in the isopod Saduria entomon, in relation to forager and resource biomasses in a system that exhibits predator–prey fluctuations in density. We found that individual diet specialization was only affected by the biomass of their preferred prey (Monoporeia affinis) and not by Saduria biomass; diet specialization was higher when Monoporeia biomass was low compared to when there were high Monoporeia biomass. Population diet breadth increased at low Monoporeia biomass whereas individual diet breadths were marginally affected by Monoporeia biomass. Overall, this led to the increase in diet specialization at low Monoporeia biomass. This study shows that predator–prey dynamics might influence diet specialization in the predator and that resource biomass, not forager biomass might be important for individual diet specialization.     

The influence of flatworm predation on zooplankton inhabiting small ponds

Experiments were performed in 1977 to determine which large zooplankton in a series of high altitude ponds can be consumed by the predatory flatworm Mesostoma ehrenbergii. This predator consumes Daphnia at a high rate and the fairy shrimp Branchinecta at a low rate, but does not consume Diaptomus. Experiments were performed in 1978 and 1979 to determined the rate of predation on Daphnia in 30 liter tubs and to determine if predation rate is correlated with surface to volume ratio of experimental containers. There is a clear correlation between surface to volume ratio and predation rate. Determinations of Mesostoma and Daphnia densities were made in a series of eight high altitude ponds, and pond surface to volume ratios were determined. Examination of these parameters lends credence to the argument that Mesostoma predation affects Daphnia dynamics in some circumstances. The results suggest that benthic invertebrate predators may affect zooplankton dynamics, especially in shallow ponds.     

Predatory behaviour ofScatophaga stercoraria under laboratory conditions

Laboratory experiments tested the effectiveness ofScatophaga stercoraria (L.) as a predator of various insect species. Photophases, ages and gender of predators and effects of dung presence on predation rates were compared for either sex using counts of prey cadavers, behavioral observations and oviposition rates. Males consistently killed more prey than females, but longer photophases did not significantly increase daily predation rates. Nine to 10-day-old flies approaching sexual maturity achieved highest predation rates. Dung was found to affect predatory behaviour but not cumulative predation. Predatory behaviour of sexually mature males was more affected than female behaviour by the presence of dung, whereas sexually immatures flies showed no response to dung presence. Preconditioning ofS. stercoraria adults using 3 prey species of different sizes did not affect the prey chosen in subsequent random choice experiments. Larger prey species such asMusca domestica (L.) orDelia antiqua (Meigen) were preferred toDrosophila sp. by both sexes ofS. stercoraria. This species may be a useful predator for future integrated pest management techniques in vegetable crops or control of house flies in barns.      

Hydroperiod,predators and the distribution of physid snails across the freshwater habitat gradient

1. Studies of species distributions across environmental gradients further our understanding of mechanisms regulating species diversity at the landscape scale. For some freshwater taxa the habitat gradient from small, shallow and temporary ponds to large, deep and permanent lakes has been shown to be an important environmental axis. Freshwater snails are key players in freshwater ecosystems, but there are no comprehensive studies of their distributions across the entire freshwater habitat gradient. Here we test the hypothesis that snail species in the family Physidae are distributed in a non‐random manner across the habitat gradient. We sampled the snails, their predators and the abiotic environment of 61 ponds and lakes, spanning a wide range in depth and hydroperiod. 2. Temporary habitats had the lowest biomass of predators. Shallow permanent ponds had the highest biomass of invertebrate predators but an intermediate fish biomass. Deep ponds and lakes had the highest fish biomass and intermediate invertebrate biomass. Five species of physids occurred in the regional species pool and 60 of the 61 ponds and lakes surveyed contained physid snails. Each pond and lake contained an average of just 1.2 physid species, illustrating limited membership in local communities and substantial among‐site heterogeneity in species composition. 3. Physids showed strong sorting along the habitat gradient, with Physa vernalis found in the shortest hydroperiod ponds and Aplexa elongata, P. gyrina, P. acuta and P. ancillaria found in habitats of successively greater permanence. When organised into a site‐by‐species incidence matrix with sites ordered according to their hydroperiods, we found the pattern of incidence to be highly coherent, showing that much of the heterogeneity in species composition from one pond to another is explained by hydroperiod. We also found that the number of species replacements along this gradient was higher than random, showing that replacement is more important than nesting in describing species composition in ponds of different hydroperiod. 4. Discriminant analysis showed that pond depth, invertebrate biomass and fish biomass were the best predictors of species composition. Analysis of these niche dimensions showed that P. vernalis and A. elongata were most successful in shallow, temporary ponds with few predators. P. gyrina and P. acuta were typically found in ponds of intermediate depth and high predator abundance. P. ancillaria was found in the deepest lakes, which had abundant fish predators but few invertebrate predators. Of the five species considered, P. ancillaria, P. vernalis and A. elongata were relatively specialised with regard to key habitat characteristics, P. gyrina was moderately generalised and P. acuta was remarkably generalised, since it alone occurred across the entire freshwater habitat gradient. The exceptional habitat breadth of P. acuta stands in contrast to distributional studies of other freshwater taxa and deserves further attention.     

Preference and consumption of Macrolophus pygmaeus preying on mixed instar assemblages of Myzus persicae

This study examines the effects of changes in the prey frequency and abundance on prey selection among the four instars of Myzus persicae by the predator Macrolophus pygmaeus under laboratory conditions. The central hypothesis was that M. pygmaeus will become more selective as prey density increases. It was also observed that M. pygmaeus can occasionally abandon a prey item that had already been killed (non-consumptive prey mortality). It was assumed that the frequency of this behavior would increase with the prey size and prey density. For these purposes prey selection was evaluated by simultaneously presenting all instars of M. persicae to the predator in equal proportions and at increasing densities. M. pygmaeus showed a higher predation rate and a higher preference for smaller prey instars at all prey densities. However, if the predation rate by the predator is expressed in terms of biomass consumed, then biomass gain was higher when feeding on the larger instars of M. persicae. The prey selectivity was indicated by the total prey mortality (consumptive plus non-consumptive prey mortality) as well as by the non-consumptive prey mortality, was associated with relatively high prey densities, depending on the prey instar. Therefore, we argued that the predatory impact of M. pygmaeus on the various instars of the aphid depends not only on prey traits but also on their relative abundance in a patch. Observed decreases in biomass gain from larger prey were likely the result of high prey availability at densities before saturation, which might have caused confusion in the predator’s prey selection.     

Partial consumption of prey: the significance of prey water loss on estimates of biomass intake

Summary A number of studies on the feeding behaviour of sucking predators have estimated the weight of biomass the predator extracts from the prey by measuring the weight change occurring in the prey. This method does not consider that a proportion of the prey weight change is lost to the immediate environment. I examined the spider Diaea sp. feeding on the fruit fly Drosophila immigrans and found that the prey lost approximately 28% more weight than the predator gained. This difference was largely explained by water loss from the prey. My results suggest that water loss, which is not available to the predator, is an important part of prey weight loss. To avoid overestimating predator biomass gain it is necessary to measure the predator weight gain directly or take into account water loss as a component of prey weight change.