Asymmetric intraspecific competition among larvae of the damselfly Ischnura elegans (Zygoptera: Coenagrionidae)

Abstract. 1. A laboratory competition experiment is described in which the growth and development rates of larvae of the damselfly Ischnura elegans (Lind.) were measured over an entire instar.
2. Two larval instars which commonly occur together in the field were used in the experiment; they were maintained with a superabundance of prey and either larvae from the same or the larger/smaller instar.
3. Small larvae suffered increased development times and decreased size increases at the moult in the presence of large larvae but similar interference effects were not evident when these smaller larvae were in the presence of other small larvae.
4. Development time and size increases of large larvae were not significantly affected by the presence of small larvae.
5. Irrespective of the instar combinations investigated, interference effects were reduced when there were more perches available, although in only a few cases was this reduction significant.
6. The consequences of the asymmetric competition reported in the experiment for the study of lifetime reproductive success in damselflies are discussed. Late emerging adults may incur reduced reproductive success.     

Stage-dependent predation on competitors: consequences for the outcome of a mosquito invasion

1. Predator-mediated coexistence occurs when predation allows competitors to coexist, due to preferential consumption of a superior competitor relative to an inferior competitor. Differences between the native treehole mosquito ( Aedes triseriatus ) and the co-occurring Asian tiger mosquito ( Aedes albopictus ) in anti-predatory larval behaviours account, in part, for the greater vulnerability of this invasive species to native predatory midge ( Corethrella appendiculata ). We test the hypothesis that stage-dependent differences in the sizes of A. albopictus and A. triseriatus larvae, relative to the size-limited C. appendiculata , contribute to differential consumption and the likelihood of predator-mediated coexistence of these competitors.
2. In all instars, larvae of A. triseriatus were larger than A. albopictus of the same stage. Third and fourth instar C. appendiculata selectively consumed late-stage A. albopictus in preference to same-stage A. triseriatus . Small, early-stage prey larvae did not differ in vulnerability to predation, but large, late-stage larvae differed significantly in vulnerability to predation, probably owing to size-limited predation by fourth instar C. appendiculata. This effect was less pronounced for third instar C. appendiculata .
3. Prey size, in conjunction with anti-predatory behavioural responses, alters the probability of predator-mediated coexistence. A stage-structured predation model showed that equally vulnerable early stages reduce the range of environmental conditions (productivities) in which predator-mediated coexistence is possible, increasing the likelihood of both competitive exclusion of the resident species or failure of the invasive to establish. These results underscore the importance of stage-dependent interspecific differences in predator–prey interactions for determining how predators may affect community composition.     

Changes in feeding behaviour of the larvae of the damselfly Enallagma cyathigerum in response to stimuli from predators

1. The feeding rates, diet composition and diel periodicity in feeding activity among larvae of the damselfly Enallagma cyathigerum Charp., exposed to chemical, visual and mechanical stimuli from the predators Ranatra linearis (L.), Notonecta glauca L. and Rutilus rutilus (L.) were investigated.
2. In response to chemical cues produced by all the predator species, larvae reduced their rate of feeding significantly (especially on large, sedentary prey).
3. Small larvae reacted more strongly than large ones to the kairomone produced by N. glauca.
4. The fish kairomone induced a reduction in feeding activity during daylight hours only. This reaction was more intense than the reaction to non-chemical cues.
5. Observed predator-induced changes in diet composition, caused directly by reduced activity during feeding, are discussed as an antipredator behaviour of damselfly larvae.     

Size-specific predation on marine invertebrate larvae

Predation on planktonic larval stages is frequently a major source of mortality for the offspring of benthic marine invertebrates. Mortality rate likely varies with larval size and developmental stage, but few experiments have measured how these factors affect predation rates. I used experimental reductions in egg size to test how variation in larval size affects the likelihood of predation during planktonic development. Blastomeres of the sand dollar Dendraster excentricus were separated at the two-cell stage to produce half-sized zygotes. Larvae resulting from this manipulation were tested for their susceptibility to predation relative to whole-sized siblings at four ages. Individuals from each size class were simultaneously presented as prey items to five predators (crab zoeae, crab megalopae, chaetognaths, solitary tunicates, and postlarval fish) in the laboratory. Four predators consumed significantly more half-sized larvae than whole-sized larvae, but one predator type (postlarval fish) consumed more whole-sized larvae. Predators that consumed more half-sized larvae also preferentially consumed younger larvae. In contrast, postlarval fish showed no significant prey preference based on larval age. These results suggest that assumptions of constant mortality rates during development should be modified to account for the effects of larval size and age.     

Further evidence for size-assortative schooling in sticklebacks

Using brook ( Culaea inconstans ) and 10-spined ( Pungitius pungitius ) sticklebacks we examined body-size related schooling behaviour. Small and large sticklebacks were allowed to choose between two test schools, of small and of large individuals, with and without a piscivorous fish ( Oncorhynchus mykiss ) visible. Sticklebacks of both species preferred the company of fish of matching body size: small fish associating with a school of small fish, large fish with a school of large fish. While no interspecific differences were found in responses to school selection, body size and predator presence did affect selection of school-type. In both species, small fish tended to show a stronger preference for matching schools. The preference was enhanced in small fish with presence of a predator, but not in large fish. These observations are further evidence for assortative schooling in sticklebacks.     

Selective predation by larval Agabus (Coleoptera: Dytiscidae) on mosquitoes: support for conservation-based mosquito suppression in constructed wetlands

1. Wetland insect predators can structure aquatic prey communities via selective predation, but receive considerably less attention than vertebrate predators. We conducted laboratory experiments to test selective predation by two species of larval dytiscid beetles ( Agabus ; Coleoptera: Dytiscidae) and the potential contribution of these beetles to suppression of mosquito populations in constructed wetlands.
2.  Agabus consumed copepods, ostracods and mosquito larvae in no-choice tests. When offered a choice, 76% of all prey consumed were mosquito larvae, indicating selective predation. Subsequent experiments revealed this preference was due to ease of capture of mosquito larvae over alternative prey.
3. Cannibalism and intraguild predation were common within and between species of Agabus , which may reduce the overall impact of the observed selective predation.
4.  Agabus larvae selectively preyed on mosquito larvae over alternative prey, which is not characteristic of some fish used as biological control agents for mosquitoes. Predator exclusion or similar experiments in the field could document how these results translate into a natural setting.
5. The findings of this study suggest developing mosquito suppression strategies focused on conservation of native wetland predators. These strategies are preferable to introducing non-native generalist predators, or applying pesticides.     

Effects of Predation Risk on Habitat Selection by Water Column Fish, Benthic Fish and Crayfish in Stream Pools

Predation risk can affect habitat selection by water column stream fish and crayfish, but little is known regarding effects of predation risk on habitat selection by benthic fish or assemblages of fish and crayfish. I used comparative studies and manipulative field experiments to determine whether, (1) habitat selection by stream fish and crayfish is affected by predation risk, and (2) benthic fish, water column fish, and crayfish differ in their habitat selection and response to predation risk. Snorkeling was used to observe fish and crayfish in, (1) unmanipulated stream pools with and without large smallmouth bass predators (Micropterus dolomieui >200 mm total length, TL) and (2) manipulated stream pools before and after addition of a single large smallmouth bass, to determine if prey size and presence of large fish predators affected habitat selection. Observations of microhabitat use were compared with microhabitat availability to determine microhabitat selection. Small fish (60–100 mm TL, except darters that were 30–100 mm TL) and crayfish (40–100 mm rostrum to telson length; TL) had significantly reduced densities in pools with large bass, whereas densities of large fish and crayfish (> 100 mm TL) did not differ significantly between pools with and without large bass. Small orangethroat darters (Etheostoma spectabile), northern crayfish (Orconectes virilis), and creek chubs (Semotilus atromaculatus) showed significantly greater densities in pools without large bass. The presence of large smallmouth bass did not significantly affect depths selected by fish and crayfish, except minnows, which were found significantly more often at medium depths when bass were present. Small minnows and large and small crayfish showed the greatest response to additions of bass to stream pools by moving away from bass locations and into shallow water. Small darters and sunfish showed an intermediate response, whereas large minnows showed no significant response to bass additions. Response to predation risk was dependent on prey size and species, with preferred prey, crayfish and small minnows, showing the greatest response. Small benthic fish, such as darters, are intermediate between small water column fish and crayfish and large water column fish in their risk of predation from large smallmouth bass.     

Risk of predation and behavioural response in three anuran species: influence of tadpole size and predator type

Many species alter their activity, microhabitat use, morphology and life history in response to predators. Predation risk is related to predator size and palatability of prey among others factors. We analyzed the predation risk of three species of tadpoles that occur in norwestern Patagonia, Argentina: Pleurodema thaul, Pleurodema bufoninum and Rhinella spinulosa. We sampled aquatic insect predators in 18 ponds to determine predator–tadpole assemblage in the study area. In laboratory conditions, we analysed the predation rate imposed by each predator on each tadpole species at different tadpole sizes. Finally, we tested whether tadpoles alter their activity in the presence of chemical and visual cues from predators. Small P. thaul and P. bufoninum tadpoles were the most vulnerable prey species, while small R. spinulosa tadpoles were only consumed by water bugs. Dragonflies and water bugs were the most dangerous tadpole predators. Small P. thaul tadpoles reduced their activity when they were exposed to all predators, while large tadpoles only reduced the activity in the presence of large predators (dragonfly larvae and water bugs). Small P. bufoninum tadpoles reduced the activity when they were exposed to beetle larvae and dragonfly larvae, while large tadpoles only reduced activity when they were exposed to larger predators (water bugs and dragonfly larvae). R. spinulosa tadpoles were the less sensitive to presence of predators, only larger tadpoles responded significantly to dragonfly larvae by reducing their activity. We conclude that behavioural responses of these anuran species were predator-specific and related to the risk imposed by each predator.     

Offspring performance in dynamic habitats: key factors for a riparian carabid beetle

Abstract.  1. Knowledge of the ecology of carabid species is largely restricted to adults, although larval mortality is assumed to be the key factor in overall mortality. As low-mobility larvae are unable to avoid unfavourable conditions, habitat selection of reproducing adults should be clearly affected by the habitat factors which determine offspring performance.
2. The present study examines the key habitat factors governing the distribution patterns of Bembidion velox larvae and adults on the river banks of the River Elbe by means of habitat suitability models. The validity of the determined habitat factors for offspring performance and survival was tested in laboratory experiments.
3. In the field, B. velox adults as well as larvae show a strong association with semi-terrestrial, sandy, open soil habitats.
4. In the laboratory, overall mortality of larvae reared in different substrates was lowest in sand of medium grain size mixed with fine and coarse sand. The first larval instars in particular reacted sensitively to variations in grain size. Furthermore, flood resistance of eggs was demonstrated, as 90% of larvae hatched under permanently flooded conditions.
5. Short development times were recorded, with 4–7 days for hatching of young larvae from eggs after oviposition and approximately 28 days for the development of adults from newly hatched larvae. This probably increases the survival probability for the pre-imaginal stages in dynamic habitats.
6. In conclusion, it can be said that reproductive success is strongly dependent on oviposition site selection by adults as this reflects the ecological demands of the immature stages.     

Oviposition habitat selection for a predator refuge and food source in a mosquito

Abstract.  1. The influence of filamentous algae on oviposition habitat selection by the mosquito Anopheles pseudopunctipennis and the consequences of oviposition decisions on the diet, development, body size, and survival of offspring were examined.
2. A natural population of An. pseudopunctipennis in Chiapas, Mexico, oviposited almost exclusively in containers with filamentous algae. Algae represented 47% of the gut contents of mosquito larvae sampled from the natural population. Mosquito larvae fed on an exclusive diet of algae developed as quickly and achieved the same adult body size (wing length) as their conspecifics fed on a standard laboratory diet.
3. Multiple regression of survival of mosquito larvae on percentage surface area cover of algae (0–99%) and the density of predatory fish (zero to four fish per container) was best described by a second-order polynomial model. Increasing fish densities resulted in a reduction in mosquito survival in all algal treatments. The highest incidence of survival was observed at intermediate (66%) algal cover in all treatments.
4. The presence of fish significantly extended larval development times whereas algal cover had no significant effect. The presence of fish resulted in emergence of smaller adults due to reduced feeding opportunities and predator avoidance behaviour. Algal cover also affected mosquito wing length but differently at each fish density.
5. Oviposition habitat selection improves survival in the presence of predators and feeding opportunities for An. pseudopunctipennis larvae.     

Tubes and tube-building in a lotic Chironomid (Diptera) community

The use of particles for the construction of tubes by a community of stream-dwelling midge (Chironomidae) larvae was investigated. It was shown that the particles present in the larval tubes occurred in the same proportions as those present on the surfaces of stones in the stream. Small particles were principally of detritus, medium-sized particles mainly of detritus and diatoms, and a a variety of types of particles, including filamentous algae and sand grains, occurred in the large size category. The absence of particle selection in the field was reflected in laboratory experiments. It is suggested that competition for suitable available space in which to build tubes and the quantity of building material are likely to be of more significance to the larval community in upland streams than particle size or type.     

Sexual size dimorphism in the two spot ladybird beetle Adalia bipunctata: developmental mechanism and its consequences for mating

Abstract  1. The literature on ladybirds indicates that males are consistently smaller than females but take the same length of time to complete their development. Rearing Adalia bipunctata at 20 and 25 °C confirmed that protandry cannot account for sexual size dimorphism in this species, nor can a difference in egg size.
2. Female larvae consumed more food and had a higher relative growth rate in the fourth instar than did male larvae.
3. When food is limited, small males appear to be more successful at mating than are large males.
4. To account for these results, it is hypothesised that the gonads of male larvae compete more strongly with the soma for resources and that this reduces the growth potential of the soma of male larvae relative to that of female larvae. The greater mating success of small males when food is limited supports the eat or mate hypothesis, which predicts that when food is limited small males will spend less time feeding and more time mating than will large males.     

Effects of zooplankton availability and foraging mode on cannibalism in three dragonfly larvae

Summary Cannibalism is likely to operate as a form of population control in dragonfly larvae. I performed aquarium experiments to investigate the effect of foraging activity and zooplankton availability on cannibalism in three dragonfly larvae. Large Cordulia aenea larvae showed low activity, and large Leucorrhinia dubia larvae showed high activity irrespective of zooplankton availability. In contrast, large Coenagrion hastulatum larvae changed from high activity in the absence to low activity in the presence of zooplankton. Small Cordulia aenea larvae were active in the absence of large conspecifics irrespective of zooplankton availability. In the presence of large conspecifics they showed a reduced activity when zooplankton were present. Small L. dubia larvae showed high activity and small Coenagrion hastulatum larvae low activity irrespective of pressence or absence of zooplankton and large conspecifics. In all three species cannibalism was highest in the absence of zooplankton. In the absence of zooplankton cannibalism was low in Coenagrion hastulatum compared to the other two species. On the contrary, in the presence of zooplankton, cannibalism did not differ between the three species.     

Evaluating the impact of predation by fish on the assemblage structure of fishes associated with seagrass (Heterozostera tasmanica) (Martens ex Ascherson) den Hartog, and unvegetated sand habitats

The role of fish predation in structuring assemblages of fish over unvegetated sand and seagrass was examined using enclosure and exclusion cages to manipulate the abundance of predatory fish from November 1998 to January 1999. In our exclusion experiment, piscivorous fish were excluded from patches of unvegetated sand and seagrass to measure how they altered abundances of small fishes, i.e., fish <10 cm in length. Habitats from which piscivorous fish were excluded contained more small fish than those with partial cages, which in turn contained more fish than uncaged areas. These patterns were consistent between unvegetated sand and seagrass areas, although the relative differences between predator treatments varied with habitat. Overall, small fish were more abundant in unvegetated sand than seagrass. Atherinids and syngnathids were the numerically dominant families of small fish and varied in complex ways amongst habitats and cage treatments. The abundance of atherinids varied inconsistently between cage treatments through time. Only during the final two sampling times did the abundance of atherinids vary significantly across cage treatments. Syngnathids were strongly associated with seagrass and were significantly more abundant in caged than uncaged habitats. In our enclosure experiment, five individuals of a single species of transient piscivorous fish, Western Australian salmon (Arripidae: Arripis truttacea Cuvier), were enclosed in cages to provide an estimate of the potential for this species to impact on small fish. The abundance of small fish varied significantly between cage treatments. Small fish were more abundant in enclosure cages and exclusion cages than uncaged areas; however, there was no difference in the abundance of small fish in enclosure cages and partial cages, and no difference between exclusion cages and partial cages. These patterns were consistent amongst habitats. Atherinids and syngnathids were again the numerically dominant families of small fish; atherinids varied more with cage structure while syngnathids did not vary statistically between cages, blocks (locations within which a single replicate of each cage treatment was applied) or habitats. Dietary analysis of caged A. truttacea demonstrated the potential for this species to influence the assemblage structure of small fish through predation - atherinids were consumed more frequently in unvegetated sand than seagrass, and syngnathids were consumed only in seagrass, where they are most abundant. Observations of significant cage or predation effects depended strongly on the time at which sampling was undertaken. In the case of the atherinids, no predation or cage effects were observed during the first two sampling times, but cage effects and predation effects strongly influenced abundances of fish during the third and fourth sampling times, respectively. Our study suggests that transient piscivorous fish may be important in structuring assemblages of small fish in seagrass and unvegetated sand, and seagrass beds may provide a refuge to fishes. But the importance of habitat complexity and predation, in relation to the potentially confounding effects of cage structure, depends strongly on the time at which treatments are sampled, and the periodicity and multiplicity of sampling should be considered in future predation studies.     

Feeding ecology of the Bay Goby (Pisces: Gobiidae). Effects of behavioral,ontogenetic, and temporal variation on diet

The ontogenetic, diel, seasonal, and yearly variations in gut fullness, diet, and prey diversity for a California estuarine gobiid (Lepidogobius lepidus Girard) were examined. Also the feeding behavior of this species was described.Small (<50 mm, SL) and large (? 50 mm, SL) gobies consumed similar prey types in different proportions. Major prey items were polychaetes, harpacticoid copepods, gammarid amphipods, molluscs, and other crustaceans. Diets of large and small gobies were not significantly correlated, and larger fish had a more diverse diet. Small fish fed at all times while larger gobies fed primarily at night. Changes in diet may be related to differential prey preferences, feeding chronologies, and increases in fish size.Both large and small gobies displayed seasonal differences in diet and prey diversity. Year-to-year changes in diet also were noted for both size classes. The bay goby uses different feeding behaviors to capture sedentary and motile prey and appears to forage opportunistically. This behavior is probably advantageous in an environment which fluctuates drastically.     

Diet and Habitat use by Bluefish, Pomatomus Saltatrix, in a Chesapeake Bay Estuary

Bluefish, Pomatomus saltatrix, are recreationally valuable finfish along the Atlantic seaboard and in the Chesapeake Bay. Diet and habitat use patterns for bluefish life history intervals in Chesapeake Bay estuaries are poorly described although it is widely acknowledged that this apex piscivorous species relies on estuarine habitat for feeding and nursery grounds after oceanic spawning and inshore migration of larvae. Bluefish diet, distribution, and abundance patterns were examined in relation to oyster reef, oyster bar, and sand bottom habitat in the Piankatank River, Virginia. Bluefish from all sites were predominantly piscivorous and were more abundant at reef sites than non-reef sites. Bluefish caught in association with the oyster reef consumed a wider diversity of prey items than fish from other sites; diet diversity may contribute to bluefish success during periods when small pelagic food fish abundance is reduced. Bluefish estuarine habitat use is positively correlated with the presence of oyster shell habitat and the complex trophic communities centering on oyster reefs.     

Interaction between insect larvae and tadpoles in tropical rain pools

Abstract. 1. Ephemeral rain pools on rock surfaces are common in Africa and are inhabited by dense populations of aquatic dipteran larvae. About 30% of the pools also support large numbers of tadpoles of the frog Ptychadena anchietae .
2. Experiments reveal that the presence of tadpoles suppress eclosion of the rock pool dwelling midge Chironomus imicola .
3. However, the presence of tadpoles also shortens the larval life-span of C. imicola by speeding up growth rates. Since the larval stages must be completed before the pool dries tadpoles may help 'fine-tune' the dipteran to the rock pool habitat.
4. A change in the diet of Cimicola larvae is associated with the accelerated development. Grazing by tadpoles results in algae, growing on the water surface, reaching the mud in tadpole faeces. These algae represent a high protein and energy food not otherwise accessible to mud dwelling dipteran larvae.
5. When pools dry tadpoles are killed, but a second species of dipteran Dasyhelea thompsoni have larvae able to survive to dry phase in situ . These larvae are scavengers so that on reflooding dead tadpoles are available as food. Experiments show that the presence of tadpoles again increase larval growth rates.
6. Both alive and dead tadpoles therefore are responsible for increasing the quality and quantity of food available to dipteran larvae and are thus among the mechanisms ensuring an abundance of food for rock pool dwelling insect larvae.     

Competition for perches between larval damselflies: the influence of perch use on feeding efficiency, growth rate and predator avoidance

SUMMARY. 1. In an investigation of perch use by two species of larval damselfies, Coenagrion puella (Linn.) and Ischnura elegans (van der Linden), larvae spaced out on a grid of vertical cocktail sticks, in the absence of food, to a greater extent than expected from a random model. Spacing was the result of direct interaction between larvae, and was not related to the level of hunger of the larvae over a 10 day starvation period.
2. Prior ownership of a perch was not a predictor of success in conflicts between larvae of the same instar. In contests between different instars larger larvae were more likely to win conflicts. Some small larvae appear to remain undetected in the presence of larger conspecifies.
3. At one level of food availability, perch ownership did not influence growth rate or mortality of C. puella or I. etegans. However, it did allow greater prey capture success rate.
4. Coenagrion puella larvae showed more movement in the absence of perches both in terms of actual distance moved and amount of swimming activity. The amount of swimming activity was reduced by addition of a predator (sticklebacks. Gasterosteus aculeatus or Pungitius pungitius) to the tank both in the absence and presence of an alternative food supply for the fish. Duration of individual swimming bouts was unaffected.
5. Sticklebacks attacked C. puella larvae significantly more often in the absence of perches, with a strong bias towards swimming larvae.
6. It is proposed that the major advantage of perch possession to C. puella larvae is in the reduction of predation as a consequence of the reduction in larval movement.     

Interspecific competition and habitat selection by the riverine dragonfly Onychogomphus uncatus

1. Substratum selection by the burrowing larvae of the dragonfly Onychogomphus uncatus was examined in an artificial laboratory stream at different larval densities and in the presence of one of three other dragonfly species.
2. The larvae of O. uncatus , as well as those of the other species, clearly preferred gravelly sand substratum rather than gravel or stone.
3. At low larval density (71.4 m^–2) in the stream, 83% of the O. uncatus were found in gravelly sand. An increase of larval abundance in the stream to 202.4 specimens m^–2 resulted in greater density in all substrata, but this increase was proportionally lowest in gravelly sand.
4. The presence of a second species had various effects on the microdistribution of O. uncatus . In the presence of Gomphus simillimus or Cordulegaster boltonii immaculifrons the distribution of O. uncatus changed significantly; their density increased in the normally less preferred substrata. This effect is interpreted as asymmetric interspecific interference. The presence of a third species, Onychogomphus forcipatus unguiculatus , had no effects.     

Influences of slimy sculpin (Cottus cognatus) predation on the rocky littoral invertebrate community in an arctic lake

We tested the role of the slimy sculpin (Cottus cognatus), a benthic fish, in structuring the rocky littoral invertebrate community in Toolik Lake, Alaska. Comparisons of sculpin gut contents and prey community structure indicated that these fish forage selectively, eating proportionally more large and motile prey, and proportionally fewer small and sessile forms. Field experiments compared the effects of natural, reduced and elevated sculpin densities on benthic community structure. At natural levels of sculpin density, biomass of trichopteran larvae were reduced by more than 50%, and predatory chironomid larvae by 27%, in comparison to areas where sculpin were excluded. Tube-dwelling and small free living chironomid larvae were unaffected at normal sculpin densities. Under artificially high sculpin densities, there was some reduction of tube-dwelling chironomids, but the small free living ones remained unaffected. There appears to be a threshold length of about 3.5 mm, below which chironomid larvae are free form sculpin predation. Tube-dwelling chironomids may be longer than this threshold, but still avoid predation by having most of their body hidden in their tubes.