Aggression and competition for shelter between a native and an introduced crayfish in Europe

1. The introduced North American crayfish, Pacifastacus leniusculus Dana, is expanding its range in Europe and locally often replaces two native crayfish species, Astacus astacus L. and Austropotamobius pallipes Lereboullet. Pacifastacus leniusculus is also expected to invade the habitat of a third native crayfish, the endangered Austropotamobius torrentium Schrank. Interspecific aggressive interactions and competition for shelter were experimentally studied in the laboratory to assess the potential impact of P. leniusculus on A. torrentium .
2. Neither species was inherently dominant in aggressive interactions, but dominance was strongly size-dependent, favouring the larger and faster growing species, P. leniusculus .
3. Access to limited shelter was generally determined by aggressive dominance, although species-specific preference also influenced the outcome of competition for shelter. Austropotamobius torrentium had a higher preference for experimental shelters and often defended these even against larger P. leniusculus .
4. In accordance with theoretical models of animal conflicts, agonistic interactions between equally sized contestants were more severe than between animals of different size.
5. The P. leniusculus used in the present experiment were infected with the crayfish plague, Aphanomyces astaci , to which the animals are resistant. The crayfish transmitted the disease to non-resistant A. torrentium which died ≈ 2 weeks after contact with P. leniusculus .     

Non‐lethal predator effects on the performance of a native and an exotic crayfish species

1. I tested the hypothesis that the potential for non‐lethal effects of predators are more important for overall performance of the fast‐growing exotic signal crayfish (Pacifastacus leniusculus Dana) than for the slower growing native noble crayfish (Astacus astacus L.). I further tested if omnivorous crayfish switched to feed on less risky food sources in the presence of predators, a behaviour that could reduce the feeding costs associated with predator avoidance. 2. In a 2 month long outdoor pool experiment, I measured behaviour, survival, cheliped loss, growth, and food consumption in juvenile noble or signal crayfish in pools with either a caged predatory dragonfly larvae (Aeshna sp.), a planktivorous fish that do not feed on crayfish (sunbleak, Leucaspius delineatus Heckel), or predator‐free controls. Crayfish had access to multiple food sources: live zooplankton, detritus and periphyton. Frozen chironomid larvae were also supplied ad libitum outside crayfish refuges, simulating food in a risky habitat. 3. Crayfish were mainly active during hours of darkness, with signal crayfish spending significantly more time outside refuges than noble crayfish. The proportion of crayfish outside refuges varied between crayfish species, time and predator treatment, with signal crayfish spending more time in refuges at night in the presence of fish. 4. Survival in noble crayfish was higher than in signal crayfish, and signal crayfish had a higher frequency of lost chelipeds, indicating a high level of intraspecific interactions. Crayfish survival was not affected by the presence of predators. 5. Gut‐contents analysis and stable isotope values of carbon (δ¹³C) and nitrogen (δ¹⁵N) indicated that the two crayfish species had similar food preferences, and that crayfish received most of their energy from feeding on invertebrates (e.g. chironomid larvae), although detritus was the most frequent food item in their guts. Signal crayfish guts were more full than those of noble crayfish, but signal crayfish in pools with fish contained significantly less food and fewer had consumed chironomids compared with predator‐free controls. Length increase of signal crayfish (35%) was significantly higher than of noble crayfish (20%), but signal crayfish in pools with fish grew less than in control pools. 6. This short‐term study indicates that fish species that do not pose a lethal threat to an organism may indirectly cause reductions in growth by affecting behaviour and feeding. This may occur even though prey are omnivorous and have access to and consume multiple food sources. These non‐lethal effects of predators are expected to be particularly important in exotic crayfish species that show a general response to fish, have high individual growth rates, and when their feeding on the most profitable food source is reduced.     

Ontogeny of microhabitat use and two-step recruitment in a specialist reef fish, the Browncheek Blenny (Chaenopsidae)

Ecological specialization is common on coral reefs and almost certainly contributes to the high diversity of fishes and invertebrates associated with reefs. Here, the recruitment pathway of an endemic Gulf of California fish, the Browncheek Blenny, Acanthemblemaria crockeri (Teleostei: Chaenopsidae), which specializes as an adult on vacant invertebrate tests or tubes, is reported. Like most reef fishes, Browncheek Blennies have a planktonic larval stage that leaves the reef and later settles on suitable habitat as a fully developed juvenile. These blennies follow a clear, “two-step” recruitment pathway, however, and do not reside in invertebrate tests until reaching an adult body size. Individual juveniles and adults were observed for 3 min intervals in order to develop average time budgets for this species. Members of both sexes and all post-settlement life-history stages were included in the analysis. The difference in habitat use by post-settlement juveniles and adults is striking; the average juvenile spends none of its time inside a test, and the average large adult spends all of its time inside a test. Using data on intermediate-sized individuals, the behavioral change associated with invading a test was determined to be size-cued, and it occurs between 20 and 30 mm standard length. Changes in feeding and predator avoidance behaviors are also associated with the ontogenetic shift from life in the open to life in a shelter. Addition of artificial shelters demonstrated the essential role of access to this specialized resource in the population regulation of adults but not juveniles of these blennies.     

Reproduction in an Irish lake population of the crayfish Austropotamobius pallipes (Lerehoullet)

SUMMARY. 1. Mature crayfish were collected in 1984 from an alkaline Irish lake before and after the mating season.
2. There was some correlation between frequency of mating and decrease in vasa deferentia weight in captive males, but in the field vasa deferentia weight losses were highly variable. The data indicate that one third of adult males may not have mated in the wild.
3. All females appear to have spawned, whether or not mated, with an average 85% reduction in ovary weights over the season.
4. Pleopodal egg counts directly after laboratory spawning were on average 81% of ovarian counts; in the field, an estimated 50 days after spawning, they were only 61% of estimated ovarian egg numbers.
5. Average ovarian egg size was positively correlated with female size. Also, larger crayfish had a wider range of egg sizes than smaller crayfish.
6. It is suggested that aggression and competition for shelter among brooding females may result in a progressive loss of eggs.     

Invasive zebra mussel colonisation of invasive crayfish: a case study

We investigated the interaction between two invasive invertebrate species in a shallow Central European flooded sandpit: the epibiosis of Ponto-Caspian zebra mussels Dreissena polymorpha on the American crayfish Orconectes limosus. Between 2004 and 2005, we followed the seasonal variation in number and size of the mussels attached to crayfish bodies, and microhabitats preferred by mussels. The proportion of crayfish colonised by mussels varied seasonally: in spring and early summer it was consistently over 75%, afterwards it dropped temporarily due to loss of bivalves during the crayfish moult, and later increased again due to re-colonisation by often relatively large juvenile mussels. Three different pathways of mussel settlement on crayfish hosts are likely: (1) primary settlement of free-swimming pediveliger larvae; (2) secondary settlement of plantigrade mussels and juveniles; (3) active re-attachment of grown mussels from the substrate to crayfish. This epibiosis was promoted by lack of suitable substrates at the studied locality. Electronic supplementary material The online version of this article (doi: ) contains supplementary material, which is available to authorized users.     

New Zealand stream crayfish: functional omnivores but trophic predators?

1. The feeding ecology of the crayfish Paranephrops planifrons in streams draining catchments in native forest and pastoral land was investigated using analyses of both stomach contents and stable isotopes of carbon and nitrogen. We aimed to (1) identify the energy sources of crayfish, (2) determine whether these were affected by ontogeny or land use change, and (3) assess the functional and trophic roles of crayfish in New Zealand hill-country streams.
2. In native forest streams, crayfish stomach contents were dominated volumetrically by leaf detritus (>60%), but in pasture streams aquatic invertebrates constituted more than 40% of stomach volumes and leaf detritus <30%. Stable isotope analyses revealed that crayfish from both native forest and pasture streams incorporated energy from aquatic invertebrates into their body tissue but did not appear to utilize detritus for growth. Therefore, deforestation had little impact on crayfish energy sources.
3. In native forest streams, adult crayfish (≥20 mm orbit-carapace length (OCL)) consumed greater amounts of leaf detritus than juvenile crayfish, but a corresponding change in stable isotope signatures was not detected. Ontogenetic shifts in diet were not consistent between land use suggesting that change in local habitat and food resources, as a result of land use change, affect crayfish food choice more than factors related solely to age or size.
4. Crayfish appear to occupy the trophic position of a predator, but by functioning as omnivores, they have dual roles as both predators and processors of organic matter. The use of gut content analysis in conjunction with stable isotope analyses revealed that the functional and trophic roles of P. planifrons differed, with implications for the interpretation of diet studies and understanding of the role of omnivores in food webs.     

Behavioural and growth differences between experienced and naïve populations of a native crayfish in the presence of invasive rusty crayfish

1. Species invasions are a leading threat to native species and ecosystems. How populations of native species respond to the presence of invasive species will ultimately determine their long-term persistence.
2. In this study, we capitalise on a unique opportunity to compare the behaviour and growth of naïve and experienced virile crayfish ( Orconectes virilis Hagen) populations in the presence of invasive rusty crayfish ( O. rusticus Girard ) . In behavioural trials, experienced O. virilis (coexisted with O. rusticus for >30 years) showed more aggressive behaviour than their naïve counterparts. Naïve O . virilis retreated from aggressive interactions with O. rusticus 38% more often than experienced O . virilis. Experienced O. virilis spent 39% more time occupying shelter than naïve O. virilis.
3. There were also differences in O. rusticus behaviour: O. rusticus spent 24% more time occupying shelter with naïve O. virilis relative to experienced O. virilis .
4. In field mesocosm experiments with O. rusticus , naïve O . virilis declined in body mass by 1% while experienced O. virilis' body mass increased by 6%, thus highlighting the potential population-level implications of the previously observed behavioural effects.
5. Our work demonstrates significant behaviour and growth differences between naïve and experienced O. virilis in the presence of invasive O. rusticus. Whether this difference is the result of phenotypic plasticity or evolution by natural selection remains to be determined. Either way, this area of inquiry has implications for managing native populations and species in an increasingly invaded world.     

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.     

Viability analysis of endangered crayfish populations

The noble crayfish Astacus astacus L. is a threatened freshwater invertebrate. Many of the remaining populations are isolated and there is considerable concern that diseases and the increased frequency of flooding events may drive these remnant populations to extinction. We performed a population viability analysis for a typical isolated noble crayfish population. We quantified the extinction risk by the mean time to extinction within 1000 years for several scenarios (flooding events, restocking of adults). For a set of parameters derived from field estimates, we estimated the mean time to extinction to be 240 years. However, the median was only 80 years. Multiple sensitivity analysis by logistic regression revealed that spawning probability, juvenile and adult mortality were the important parameters for the survival of the population. The mean time to extinction decreased with increasing frequency of floodings. This is alarming, considering the magnitude of the effect and the expectation of an increasing number of floodings with global warming. Restocking, however, was found to have only a minor effect on the mean time to extinction. Overall, our simulations suggested that for the long-term and self-sustaining survival of the noble crayfish, particularly where they remain isolated, we have to improve the extent and quality of habitats. Nevertheless, additional measures are necessary, especially the removal of dispersal barriers to allow some exchange of individuals between populations. However, this also calls for a control of invasive crayfish species.     

Managing invasive crayfish: use of X-ray sterilisation of males

1. In the last decade, many attempts have been made to contain the spread of the North American crayfish Procambarus clarkii , an invader of many European water bodies, but none has been successful.
2. This study investigates the effects that ionising irradiation has on the reproduction of P. clarkii males by analysing the behaviour of the treated individuals, their reproductive output, the vitality and survival of their offspring and the damages induced to their gonads.
3. We found that a dose of 20 Gy X-rays did not compromise either the survival or the mating ability of males, but reduced the size of their testes and significantly altered spermatogenesis. The number of aborted eggs was larger in clutches sired by treated males, with a consequent reduction (by 43%) in the number of offspring.
4. These results foster hopes in our efforts to control invasive crayfish, showing the potential effectiveness of ionising irradiation to reduce male fertility in the perspective of adopting the Sterile Male Release Technique to control P. clarkii populations.     

The artificial incubation of crayfish eggs: review and report from an experimental study concerning the effects of offspring origin (maternal or artificial incubation) on the survival and growth of juvenile signal crayfish (Pacifastacus leniusculus, Astacidae)

The development of artificial incubation techniques in astacid crayfish has attracted attention from scientists in many countries ever since the nineteenth century. It is only in the last few years that these techniques, along with studies on egg storage and transport, have provided reliable options for improving the reproductive phase in farming. The juveniles produced need to be reared until they reach a sufficient size both for restocking and for growing purposes. In view of the current level of knowledge of rearing juvenile astacids, two 80-day experiments were carried out under controlled conditions to compare the survival and growth of Stage 2 juvenile signal crayfish (Pacifastacus leniusculus) from two origins: maternal or artificial incubation. In the first experiment, three treatments were tested: juveniles from artificially incubated eggs with formaldehyde treatments, juveniles from maternal incubation and a mixture from both origins (50% each). Survival rates ranged from 87.8% to 93.3% with no significant differences among treatments. Crayfish from artificial incubation grew significantly faster (11.47 mm carapace length (CL), 373.80 mg weight) than crayfish from maternal incubation. In the second experiment, a bifactorial design included four treatments: the crayfish was derived from artificial or from maternal incubation and was fed once a day or twice a day. Final survival rates ranged from 68.89% to 77.78%, with no significant differences among treatments. Crayfish from artificial incubation grew significantly faster than crayfish from maternal incubation. The highest CL (14.54 mm) and weight (780.13 mg) were reached by the juveniles from artificial incubation that were fed once a day. No significant differences were found between the two feeding frequencies. Results showed that artificial incubation with formaldehyde treatments had no harmful effects and made it feasible to get a better performance from the juveniles obtained.     

A laboratory study of the effects of shelter availability and invasive crayfish on the growth of native stream fish

1. The alteration of stream habitats by urbanisation reduces the availability of shelter. Reduced shelter availability may increase both predation risks and metabolic costs, negatively affecting the growth performance of stream fish. Although urbanised streams often allow the establishment of invasive species, the additive or interactive effects of shelter availability and invasive species are rarely explored. The invasive red swamp crayfish (Procambarus clarkii) is a strong shelter competitor and predator in streams. 2. We conducted a stream‐channel experiment to investigate how shelter availability and this invasive crayfish species affect the growth performance (measured as the loss in body mass under starved conditions) of two native benthic species, Japanese crucian carp (Carassius auratus complex) and an endangered cobitid fish (Lefua echigonia). 3. We found that both the absence of shelter and the presence of crayfish increased body mass loss in the crucian carp. However, the growth performance of the crucian carp was improved by the presence of shelter except in the presence of crayfish, in which case the shelter available did not ameliorate the negative effect of the crayfish on growth performance (i.e. an interactive effect). This result suggests that crayfish may affect the growth performance of the crucian carp through chemical cues as well as through shelter occupancy. 4. In contrast, shelter availability is the primary factor affecting the body mass loss of the cobitid fish. However, the growth performance of these fish was unaffected by the presence of crayfish. The change in the growth performance of the cobitid fish in response to the absence of shelter was greater than that of the crucian carp. This finding suggests that the cobitid fish may be more vulnerable than the crucian carp to a reduction in shelter availability. 5. Our study demonstrates that reduced shelter availability and/or crayfish invasion can have significant, indirect negative effects on the growth performance of native fish, but whether those effects are interactive might vary depending on the species. Our findings have clear implications for stream restoration and habitat assessment.     

Bioenergetics and phenology of reproduction of the introduced red swamp crayfish, Procambarus clarkii, in Do·ana National Park, Spain, and implications for species management

1. The red swamp crayfish, Procambarus clarkii (Girard), is a North American species which was introduced into the Lower Guadalquivir, Spain, in 1974. Procambarus clarkii has become a key species since it has greatly altered the functioning and structure of the ecosystems it has invaded.
2. Since P. clarkii spawn spontaneously throughout the year, it is to be expected that their ecological impact on the ecosystem may vary depending of the timing of habitat flooding (e.g. temporary freshwater marsh, and temporary or permanent pond). The objective of the present study was to understand the reproductive phenology and bioenergetics of P. clarkii in Do·ana National Park, Spain, freshwater ecosystems, and consequently, propose some management plans for their populations.
3. Five stages of ovarian development were detected. Final ovarian maturation occurred a few days before spawning and spawning was synchronized among mature females of all populations. Egg numbers hatched in pleopods ranged from 450 to 550. Timing of juvenile release from pleopods varied greatly (10–35 days).
4. Gravid females support most of their metabolic needs by catabolizing carbohydrates (overall mean of O:N = 30.39), and to a lesser extent, lipids. The aerobic and excretory metabolisms of gravid females were lower than those of immature animals of similar weight.
5. Analysis suggested that several factors influence the spawning of crayfish at the monitored sites. The major factors are water regime and temperature. Red swamp crayfish appear to be a spontaneous spawning species which may adjust their reproductive cycle to the timing of flooding, enhancing the utilization of available resources.
6. Populations of P. clarkii in Do·ana National Park could be controlled by the management of the water regime and by devising a fishing scheme which would maintain an appropriate population structure.     

Habitat complexity reduces aggression and enhances consumption in crayfish

We tested the hypothesis that increased habitat complexity would reduce intraspecific interactions among crayfish (Orconectes propinquus), and result in an increase in the consumption rate of prey at different crayfish densities. The effect of crayfish density, food level (trout eggs), and habitat complexity on prey consumption by crayfish was quantified in the laboratory. There was a significant difference in the consumption rate between different food levels. When food was scarce, almost all trout eggs were consumed regardless of crayfish density or habitat complexity. When food was unlimited, there was a significant positive linear relationship between eggs consumed per crayfish and habitat complexity. However, the relationship was not significant when trials without habitat were deleted from the analysis. We found that habitat complexity significantly reduced intraspecific aggression. Our findings suggest that a minimal amount of habitat complexity can reduce interactions among predators, ultimately resulting in increased prey consumption.     

Effects of Adult Dreissena polymorpha on Settling Juveniles and Associated Macroinvertebrates

The impact of Dreissena polymorpha settlement on recruitment of juvenile mussels and density of other macroinvertebrates was studied in field experiments using blank concrete blocks and tiles (control), blocks and tiles with attached empty zebra mussel shells, and blocks and tiles with attached living mussels. On blocks, dominant invertebrate taxa showed colonization patterns coinciding with increased habitat complexity owing to zebra mussel settlement or the biodeposition of faeces and pseudofaeces. Adult and especially juvenile zebra mussels preferred blocks with empty shells to blank blocks and blocks with living mussels; this might possibly be caused by a chemical cue that induces gregarious settlement. Lower recruitment on blocks with attached living mussels compared to blocks with only shells could be the consequence of ingestion of larvae by adult mussels and of competition for food. On tiles, the sediments deposited and the organic content of the sediment were investigated. Sedimentation was significantly higher on shell‐only and live‐mussel tiles compared to blank tiles. Organic matter differed significantly between blank and live‐mussel tiles.     

Adult Male Density Influences Juvenile Microhabitat Use in a Territorial Lizard

Habitat choice often has strong effects on performance and fitness. For many animals, optimal habitats differ across age or size classes, and individuals shift habitat use through ontogeny. Although many studies document ontogenetic habitat shifts for various taxa, most are observational and do not identify the causal factor of size‐specific habitat variation. Field observations of the brown anole lizard (Anolis sagrei) show that juveniles perch on shorter and thinner vegetation than adults. We hypothesized that this variation is due to adult males forcing smaller juveniles to less preferred habitat. To test this assertion, we manipulated adult male densities in mesh enclosures with artificial trees to examine the response of juvenile microhabitat choice. We found that adult male density had strong effects on juvenile perch height, perch width, and substrate use, suggesting that age‐class competition contributes to the observed ontogenetic differences in habitat choice. We also found that time of day significantly affected juvenile perch height and substrate use. In many cases, our results suggest that juveniles distance themselves from adults using different microhabitats from those used in our control ‘no‐adult’ treatment. However, these findings were often body size dependent and varied depending upon time of day. This study highlights the complexity of juvenile perching behavior and demonstrates the role of intraspecific interactions in shaping habitat use by juvenile animals.     

A minimalist approach to the effects of density-dependent competition on insect life-history traits

Abstract  1. Due to its effects on the phenotypic and genotypic expression of life-history traits, density-dependent competition is an important factor regulating the growth of populations. Specifically for insects, density-dependent competition among juveniles is often associated with increased juvenile mortality, delayed maturity, and reduced adult size.
2. The aim of the work reported here was to test whether the established phenotypic effects of density-dependent competition on life-history traits could be reproduced in an experimental design requiring a minimal number of individuals. Larvae of the mosquito Aedes aegypti were reared at densities of one, two, or three individuals per standard Drosophila vial and in six different conditions of larval food availability. This design required relatively few individuals per independent replicate and included a control treatment where individuals reared at a density of one larva per vial experienced no density-dependent interactions with other larvae.
3. Increased larval densities or reduced food availability led to increased larval mortality, delayed pupation, and the emergence of smaller adults that starved to death in a shorter time (indicating emergence with fewer nutritional reserves).
4. Female mosquitoes were relatively larger than males (as measured by wing length) but males tended to survive for longer. These differences increased as larval food availability increased, indicating the relative importance of these two traits for the fitness of each sex. The role of nutritional reserves for the reproductive success of males was highlighted in particular.
5. This minimalist approach may provide a useful model for investigating the effects of density-dependent competition on insect life-history traits.     

Benefits of adjacent habitat patches to the distribution of a crayfish population in a hydro-dynamic wetland landscape

Aquatic macrophyte patches are natural features of wetland ecosystems that serve as habitat for aquatic animals. Previous studies suggest animal densities in Everglades, USA, wetlands are generally less numerous in sawgrass ridges than in deeper lily sloughs. We studied the density distribution of a population of Procambarus fallax in ridge and slough habitat types over a 2-year period, spanning two wet–dry cycles and estimated growth and survival rates under flooded conditions to understand comparative value of each to the crayfish population. Procambarus fallax individuals inhabited and recruited in both marsh habitats. During periods of high water, crayfish densities were similar in both habitats; however, densities in both habitats varied seasonally, leading us to postulate some degree of population redistribution in response to fluctuating water depths. Analysis of size distributions over time revealed juveniles in both habitats and two major recruitment periods each year; distinct juvenile cohorts were present in early winter (Nov–Dec) and mid-summer (July–Aug). An in situ experiment of juvenile growth demonstrated that slough habitat type supported faster growth over ridge habitat. To understand habitat-specific mortality risk, a tethering study during flooded conditions indicated that relative predation risk by aquatic predators was greater in sloughs for all sizes and higher for smaller individuals in both habitats. The comparative importance of ridge and slough balances growth potential and survival probability during flooded conditions. This is the first study through time and across both habitat types analyzing the distribution and size structure of P. fallax population in the Everglades.     

Diet of the leeches Erpobdella octoculata (L) and Helobdella stagnalis (L) in a lotic habitat subject to organic pollution

SUMMARY 1. A study of the diet of E. octoculata (L) and H. stagnalis (L) using a serological technique was carried out in a lot ic habitat subject to organic pollution and seasonal sewage fungus growth.
2. The results indicate that E. octoculata consumed predominantly Chironomidae and Oligochaeta, while H. stagttalis had a less pronounced prey preference, consuming rather similar proportions of chironomids, oligochaetes, Asellus and gastropod snails.
3. At enriched sites, E. ocioculata consumed relatively more chironomids and fewer oligochaetes, while H. stagnalis took more gastropods and fewer chironomids.
4. Seasonal differences were noted in the diet of both leeches with chironomids predominating chiefly from April to August, and molluscs, oligochaetes and Asellus at other times of the year. Intra-specific differences (on a weight-age basis) were only seen for E. octoculata , small forms consuming chiefly oligochaetes and larger ones concentrating on chironomids.     

Feeding ecology of larval and juvenile American shad (Alosa sapidissima) in a small pond

Release ponds are used as part of a multifacet effort to restore American shad ( Alosa sapidissima ) in the upper Susquehanna River basin. Little information exists, however, on the feeding ecology of young shad in small ponds. Consequently, we examined feeding ecology and prey selection of 299 larval and 299 juvenile American shad in a small pond during spring and summer. Larval shad mainly consumed copepods (37.7%) and cladocerans (37.4%) whereas juvenile shad ate chironomids (43.1%) and ostracods (28.4%). Larval and juvenile shad exhibited diel variation in diet composition and feeding periodicity. Food consumption by shad was minimal at night; feeding activity was highest during the day, peaking at 2000 h for both larvae and juveniles. Electivity values of shad larvae for prey taxa were highest for cladocerans (+0.27) and lowest for ostracods (−0.07). Electivity values of juvenile shad were highest for chironomids (+ 0.21) and ostracods (+ 0.09), and lowest for copepods (− 0.08) and baetids (− 0.14). Our data indicate differences in diet composition, prey preference and, to a lesser extent, feeding patterns between larval and juvenile American shad in small pond environments.