Coupling long-term studies with meta-analysis to investigate impacts of non-native crayfish on zoobenthic communities

1. Biological invasions are widely recognised as a significant component of human‐caused environmental change and a primary threat to native biodiversity. The negative impacts of species invasions are particularly evident for freshwater crayfish faunas. 2. This study provides novel insight into the ecological effects of native and non‐native crayfish on zoobenthic communities (with emphasis on the non‐native rusty crayfish, Orconectes rusticus) across broad scales by combining a meta‐analysis of small‐scale experimental studies with a long‐term observational study conducted over a 24 year period in Sparkling Lake, Wisconsin, U.S.A. (46°00′N, 89°42′W). 3. The meta‐analysis summarised quantitatively the results of cage experiments for seven species of crayfish spanning four continents. We found that total zoobenthos densities (primarily Gastropoda and Diptera) were significantly lower in treatments containing crayfish relative to controls; a result that was significant for non‐native crayfish but not for crayfish in their native range, perhaps owing to a small sample size. In contrast to other species, rusty crayfish were also negatively associated with Ephemeroptera. 4. Results from the time series analysis comparing temporal trends in rusty crayfish and invertebrate abundances from Sparkling Lake were consistent with the findings from the meta‐analysis. Rusty crayfish were negatively correlated with the abundance of total zoobenthos, Diptera, Ephemeroptera and Odonata, as well as families of Trichoptera. 5. By coupling the results from short and long‐term research, our study offers greater insight into the nature of crayfish‐invertebrate interactions in aquatic systems, revealing consistent effects of invasive crayfish on native fauna. The control and management of invasive species is facilitated by the knowledge that well executed small‐scale studies may be extrapolated to understand larger‐scale ecological interactions.     

Effects of an invasive crayfish on trophic relationships in north‐temperate lake food webs

1. The introduction of invasive species is one of the main threats to global biodiversity, ecosystem structure and ecosystem processes. In freshwaters, invasive crayfish alter macroinvertebrate community structure and destroy macrophyte beds. There is limited knowledge on how such invasive species‐driven changes affect consumers at higher trophic levels. 2. In this study, we explore how the invasive rusty crayfish Orconectes rusticus, a benthic omnivore, affects benthic macroinvertebrates, as well as the broader consequences for ecosystem‐level trophic flows in terms of fish benthivory and trophic position (TP). We expected crayfish to decrease abundance of benthic macroinvertebrates, making most fish species less reliant on benthic resources. We expected crayfish specialists (e.g. Lepomis sp. and Micropterus sp.) to increase their benthic dependence. 3. In 10 northern Wisconsin lakes, we measured rusty crayfish relative abundance (catch per unit effort, CPUE), macroinvertebrate abundance, and C and N stable isotope ratios of 11 littoral fish species. We used stable isotope data and mixing models to characterise the trophic pathways supporting each fish species, and related trophic structure to crayfish relative abundance, fish body size and abiotic predictors using hierarchical Bayesian models. 4. Benthic invertebrate abundance was negatively correlated with rusty crayfish relative abundance. Fish benthivory increased with crayfish CPUE for all 11 fish species; posterior probabilities of a positive effect were >95%. TP also increased slightly with crayfish CPUE for some species, particularly smallmouth bass, largemouth bass, rock bass and Johnny darter. Moreover, both fish body size and lake abiotic variables explained variation in TP, while their effects on benthivory were small. 5. Rusty crayfish abundance explained relatively little of the overall variation in fish benthivory and TP. Although rusty crayfish appear to have strong effects on abundances of benthic macroinvertebrates, energy flow pathways and trophic niches of lentic fishes were not strongly influenced by invasive rusty crayfish.     

Intraguild predation between spawning smallmouth bass (Micropterus dolomieu) and nest‐raiding crayfish (Orconectes rusticus): implications for bass nesting success

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Testing the differential predation hypothesis for the invasion of rusty crayfish in a stream community: laboratory and field experiments

1. We tested the hypothesis that the non‐native rusty crayfish (Orconectes rusticus) is less vulnerable to predators than two native species (O. propinquus and O. obscurus) it is replacing in streams of the upper Susquehanna River catchment (New York, U.S.A.). 2. We used laboratory experiments to compare species‐specific predation rates by smallmouth bass (Micropterus dolomieu) on crayfish of equal size and field tethering experiments to compare relative predation rates between native O. propinquus and non‐native O. rusticus by the suite of crayfish predators in our system. We predicted that crayfish size would affect predation rate but that predation rates would be equal among species when size was controlled. 3. We also tested for two potential artefacts of tethering. We tethered crayfish in cages to test whether the ability to escape from tethers is size specific, and we tested whether tethering alters differential predation among crayfish species by the smallmouth bass. 4. In the laboratory, smallmouth bass predation on rusty crayfish was lower than on either of the native species. In the field, predation risk for tethered crayfish was inversely related to size but did not differ among species when size was taken into account. Because rusty crayfish in the field experiment were slightly larger than the native species, as in nature, mortality was overall lower for the rusty crayfish. 5. In cages, smaller crayfish were more probably to escape from tethers than larger ones, an artefact that may partially confound results from our tethering experiments. Unexpectedly, tethering nearly eliminated predation by smallmouth bass. This artefact prevented us from testing for an interaction of tethering with differential predation and means that the results of field tethering experiments do not include any contribution from smallmouth bass predation. 6. Our experiments highlight the importance of explicitly considering potential artefacts that could confound results. 7. Our results indicate that differential predation contributes to the rusty crayfish's invasion of a stream community. In our study system, predation rates on rusty crayfish are lower than for native species mostly because of selection by predators for smaller crayfish; species‐specific characteristics such as behaviour that further reduce predation may also contribute, especially where smallmouth bass predation is important.     

Predation on invasive zebra mussel, Dreissena polymorpha, by pumpkinseed sunfish, rusty crayfish, and round goby

The enemy release hypothesis states that invasive species are successful in their new environment because native species are not adapted to utilize the invasive. If true for predators, native predators should have lower feeding rates on the invasive species than a predator from the native range of the invasive species. We tested this hypothesis for zebra mussel (Dreissena polymorpha) by comparing handling time and predation rate on zebra mussels in the laboratory by two North American species (pumpkinseed, Lepomis gibbosus, and rusty crayfish, Orconectes rusticus) and one predator with a long evolutionary history with zebra mussels (round goby, Neogobius melanostomus). Handling time per mussel (7 mm shell length) ranged from 25 to >70 s for the three predator species. Feeding rates on attached zebra mussels were higher for round goby than the two native predators. Medium and large gobies consumed 50–67 zebra mussels attached to stones in 24 h, whereas pumpkinseed and rusty crayfish consumed <11. This supports the hypothesis that the rapid spread of zebra mussels in North America was facilitated by low predation rates from the existing native predators. At these predation rates and realistic goby abundance estimates, round goby could affect zebra mussel abundance in some lakes.     

Habitat, predation, and coexistence between invasive and native crayfishes: prioritizing lakes for invasion prevention

Conditions fostering coexistence of native species with invasive species have received little attention in invasion biology, especially for closely related invasive and native species. We used long-term datasets on multiple replicate invasions to define conditions under which native virile crayfish (Orconectes virilis) can coexist with invasive rusty crayfish (O. rusticus). We examined multiple drivers of coexistence involving habitat use and predation at between-lake and within-lake scales to derive predictions that could guide prioritization efforts to prevent future introductions of rusty crayfish and mitigate impacts of existing invasions. Lakes in which native species persisted for many years had significantly less cobble and sand habitats, and significantly more vegetated habitats compared to lakes from which native crayfish have been displaced. In the presence of rusty crayfish, virile crayfish alter their habitat use to vegetated habitats relative to habitat use in the absence of rusty crayfish. Such vegetated habitats had greater plant standing crop, plant species richness, and sediment percent organic matter compared to vegetated sites occupied by rusty crayfish. Our results suggest that low abundance of cobble habitat and altered habitat use allows native crayfish to coexist with the rusty crayfish invader. At the within-lake scale, virile crayfish persist by escaping predation in the vegetated habitats, despite suboptimal abiotic conditions. By understanding these abiotic and biotic conditions that promote coexistence, managers could enhance native crayfish persistence by targeting high cobble lakes for efforts to prevent the introduction of invasive crayfish, and targeting vegetated habitats for protection in already invaded lakes.     

An analysis of the mechanisms governing species replacements in crayfish

Summary We investigated mechanisms governing replacement of the native crayfish Orconectes sanborni by an invading cryafish, Orconectes rusticus. The two species had similar life histories, habitat preferences, and feeding patterns in allopatric and sympatric stream areas. Orconectes rusticus young-of-year (YOY) grew faster than O. sanborni YOY in the field. Adult O. rusticus were larger and, hence, dominant over adult O. sanborni; YOY were non-aggressive. In laboratory experiments, adult crayfish (about 28 mm carapace length or larger) were not susceptable to predation by largemouth bass (Micropterus salmoides, 30 cm total length) and did not alter shelter use when fish were present. Orconectes rusticus YOY were less susceptible to predation than O. sanborni YOY. Orconectes rusticus YOY reduced their vulnerability to largemouth bass by occupying shelters more often than YOY O. sanborni. In mixed-species mateselection experiments, male O. rusticus and male O. sanborni preferentially mated with O. rusticus females. Inappropriate mate selection in sympatry may have caused the 90% reduction in recruitment for both species in 1982. Orconectes rusticus probably maintains greater population growth than O. sanborni, because (1) more gravid O. rusticus females occurred in sympatry, (2) O. rusticus produced more young than O. sanborni, and (3) O. rusticus young grew faster. Reproductive interference, acting synergistically with differences in aggressive dominance and young-of-year susceptibility to predation, appears to serve as the major mechanisms regulating replacement of O. sanborni by O. rusticus in Ohio streams.     

Reciprocal Relationships Between Exotic Rusty Crayfish,Macrophytes, and <Emphasis Type="Italic">Lepomis</Emphasis> Species in Northern Wisconsin Lakes

Abstract The non-native rusty crayfish (Orconectes rusticus) has invaded many lakes of northern Wisconsin, profoundly changing littoral zones in the process. There are other lakes that have been invaded, but do not exhibit these changes. We hypothesized that endogenous feedbacks could form involving rusty crayfish, the macrophytes they destroy, and Lepomis species whose abundance is positively related to macrophyte abundance and also consume juvenile crayfish. We assessed this proposal with long-term data from one lake, a regional comparative study, and a case study of Lepomis predation on crayfish. Through time and across lakes, abundances of rusty crayfish, littoral macrophytes and species of the genus Lepomis were related in a fashion that indicated a set of feedbacks that regulate the abundance of all three. Intense predation on juvenile crayfish by abundant Lepomis is capable of maintaining some crayfish populations at low abundance. Thus, some lakes display profound ecological changes where crayfish achieve high abundance, and others sustain crayfish at low abundance. Consequently, lakes invaded by rusty crayfish may take on the appearance of alternative ecological regimes. Direct experimentation is necessary to determine if, and under what conditions, a lake can exist in either regime. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Infection with a trematode parasite differentially alters competitive interactions and antipredator behaviour in native and invasive crayfish

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The rapid spread of rusty crayfish (Orconectes rusticus) with observations on native crayfish declines in Wisconsin (U.S.A.) over the past 130 years

The rusty crayfish, Orconectes rusticus, is one of America’s best-known non-indigenous crayfishes, having been identified as extirpating native crayfishes and disrupting local aquatic ecosystems. Over the past 40–50 years, rusty crayfish have spread from its historical range in the Ohio River drainage (U.S.A.), to waters throughout much of Illinois, Michigan, Wisconsin, and Minnesota and parts of 11 other states, Ontario (Canada) and the Laurentian Great Lakes. Using a comprehensive dataset based on all known historical records and extensive present-day surveys (n = 2775) this study reports on the invasion history of rusty crayfish, with observations on concomitant declines of native crayfishes in Wisconsin over the past 130 years (1870–2004). We found that rusty crayfish occurrences have increased from 7% of all crayfish records collected during the first 20 years of their invasion (1965–1984) to 36% of all records during the last 20 years, and that rusty crayfish have replaced the northern clearwater crayfish (O. propinquus) and virile crayfish (O. virilis) as the most dominant member of the contemporary crayfish fauna. In light of our results we discuss the introduction, establishment and integration phases of the rusty crayfish invasion and provide preliminary predictions of the potential distribution of rusty crayfish in Wisconsin lakes based on critical environmental requirements.     

A trematode parasite alters growth,feeding behavior,and demographic success of invasive rusty crayfish (Orconectes rusticus)

Nonindigenous species can cause major changes to community interactions and ecosystem processes. The strong impacts of these species are often attributed to their high demographic success. While the importance of enemy release in facilitating invasions has often been emphasized, few studies have addressed the role of parasites in the invasive range in controlling demographic success of potential invaders. Here we examine whether a trematode parasite (Microphallus spp.) can contribute to previously documented alternate states in the abundance of invasive rusty crayfish (Orconectes rusticus) in north temperate lakes in Wisconsin, USA. Microphallus infect O. rusticus after emerging from their first intermediate host, a hydrobiid snail. As previously documented, O. rusticus reduce densities of hydrobiid snails through direct predation and destruction of macrophyte habitat. Therefore, if Microphallus substantially reduce O. rusticus fitness, these parasites may reinforce a state of low crayfish abundance, and, at the other extreme, abundant crayfish may repress these parasites, reinforcing a state of high crayfish abundance. From samples collected from 109 sites in 16 lakes, we discovered (1) a positive relationship between crayfish infection intensity and hydrobiid snail abundance, (2) a negative relationship between parasite prevalence and crayfish abundance, and (3) a negative relationship between parasite prevalence and crayfish population growth. With experiments, we found that infection with Microphallus reduced foraging behavior and growth in O. rusticus, which may be the mechanisms responsible for the population reductions we observed. Overall results are consistent with the hypothesis that Microphallus contributes to alternate states in the abundance and impacts of O. rusticus.     

Preemptive legislation inhibits the anthropogenic spread of an aquatic invasive species, the rusty crayfish (Orconectes rusticus)

The enactment of legal policies is often recommended to prevent anthropogenic introductions of invasive species. In this paper, we evaluated the effectiveness of proactive state legislative policies in deterring colonization by rusty crayfish (Orconectes rusticus) and the expected spread into previously uninvaded states using network-based spatial analysis. We found that the presence of regulations was positively associated with the presence of rusty crayfish (p = 0.027), but often regulations were put into effect subsequent to the invasion. Regulations that did not explicitly prohibit transport, applied to specific drainages, or prohibited only rusty crayfish were not effective. However, preemptive legislation was effective in reducing the likelihood of invasion, if regulations prohibited the transport of all live crayfish species between water bodies, as only 1 state which passed such legislation prior to invasion by rusty crayfish was subsequently colonized (S = 12, p = 0.031). Five states are likely to be invaded by rusty crayfish via range expansion across interstate drainages, and dispersal rates suggest that all of these states could be colonized within 10 years. While it is unlikely that regulations will prevent dispersal-based invasion across state lines through shared drainages, preemptive legislation can significantly reduce anthropogenic spread of aquatic invasive species between drainages and effectively retard the expansion of rusty crayfish. Our results suggest that the most effective form of legislation is one that does not require individuals to identify species, thus we recommend states enact policies that explicitly prohibit the transport of all live crayfish between water bodies.     

The influence of evolution and plasticity on the behavior of an invasive crayfish

Invasion success can be enhanced by evolution and behavioral plasticity, but the importance of these processes for most invasions is not well understood. Previous research suggests there is a genetic basis for differences in growth rate between native and invaded range rusty crayfish (Orconectes rusticus). We hypothesized that invaded range O. rusticus achieve faster growth by allocating more time to foraging and less to defense. We conducted a laboratory experiment to test the effects of range (native or invaded) and plasticity (as induced by exposure to predators) on crayfish behavior. We collected O. rusticus adults and eggs from both ranges, hatched eggs in the lab, and reared juveniles in common conditions either with or without predatory fish. We then quantified adult and juvenile crayfish activity in an experiment with and without predatory fish. In support of our hypothesis, invaded range adults displayed reduced antipredator behavior compared to native range adults. Further, invaded range juveniles were more active than native range juveniles without predators, but all juveniles were inactive with predators. In addition, invaded range juveniles had greater plasticity in behavior than native range juveniles. These results suggest that activity level in the absence of predators has diverged in the invaded range. Because active crayfish consume more prey, this change in behavior may be responsible for rapid growth in the invaded range of O. rusticus, a trait that contributes to the strong ecological impacts of this invasive crayfish.     

A Test of Temporal Variation in Risk and Food Stimuli on Behavioral Tradeoffs in the Rusty Crayfish, Orconectes rusticus: Risk Allocation and Stimulus Degradation

The effects of temporal variation in exposure to predation risk on behavioral tradeoffs were tested in the rusty crayfish, Orconectes rusticus. Based on the risk allocation hypothesis, we predicted that increasing the frequency of encounter with predation risk would yield increasing responses to a food stimulus in the presence of both a risk stimulus and a food stimulus. Crayfish were exposed to risk every 12 h, every 6 h, or left undisturbed for 24 h prior to testing. The risk stimuli used were a plain water control, snapping turtle (Chelydra serpentina) cue, and conspecific alarm cue. After 24 h of conditioning, the crayfish were exposed to a combination of risk cue and food cue. The behavioral responses of the crayfish were recorded for 5 min immediately following the introduction of the cues and again for 5 min, 1 h after stimulus exposure. The crayfish were observed at the two times to determine how their responses to the combination of risk and food cues changed over time. The responses of the crayfish were significantly influenced by stimulus treatment, time, and the interaction of time and stimulus treatment. Further analysis indicated that responses to the stimulus treatments changed differently over time. Immediately after exposure, the crayfish were more active in the control and snapping turtle treatments than in the conspecific alarm treatment. The high levels of activity initially observed in the control and snapping turtle treatments waned over time, such that the behaviors recorded 1 h after exposure were not significantly affected by stimulus treatment. Neither frequency nor the interactions of frequency with stimulus and/or time significantly affected crayfish behavior. The results of this study did not support the risk allocation model and contrast with results from similar work on the virile crayfish, Orconectes virilis.     

Invasion of an occupied niche by the crayfish Orconectes rusticus: potential importance of growth and mortality

We are exploring mechanisms of an invasion that contradicts the oft-cited generalization that species invade vacant niches. In northern Wisconsin lakes, the introduced crayfish Orconectes rusticus is replacing two ecologically similar resident congeners, O. virilis and O. propinquus. In laboratory experiments, we compared growth and mortality of individually maintained crayfish offered one of five ad libitum diets: invertebrates, macrophytes, dentritus, periphyton or all items combined. Mortality was highest for O. virilis and lowest for O. rusticus. Macrophyte diets yielded the highest mortality. All three species grew best on invertebrate and combination diets but grew little or not at all on diets of periphyton, detritus or macrophytes. O. rusticus and O. virilis grew more than O. propinquus. O. rusticus grew more quickly and/or was better able to survive overall than its congeners. Therefore, O. rusticus would probably have advantages over O. virilis and O. propinquus in competitive interactions, reproductive success and avoiding size-selective fish predation. Subtle interspecific differences may interact strongly with other ecological factors and contribute to the displacement of resident species from a well-occupied niche.     

The impacts of flow on chemical communication strategies and fight dynamics of crayfish

Signal transmission is influenced by the physics of an environment. Consequently, a physical effect on sensory signals can influence how animals send or sample sensory information. Habitat-specific physics may constrain or enhance signal transmission (e.g. sound transmission in a flowing river versus a still pond) and provide a mechanism for the evolution of sensory biases. This study investigated how the transmission of chemically mediated social signals in crayfish is influenced by two different aquatic environments. Agonistic bouts between crayfish were performed under lotic (flowing water) and lentic (nonflowing, still water) conditions. When crayfish (Orconectes rusticus) collected from a lotic system (river) interacted under lotic conditions, we noted that dominant O. rusticus spent more time upstream than subordinate O. rusticus. Orconectes rusticus positioned themselves randomly and spent equal amounts of time with respect to upstream and downstream in the nonflowing environment. We tested another species, Orconectes virilis, collected from a nonflowing environment (lake) and they showed no positional preference when tested in flow. Additionally, both O. rusticus and O. virilis took longer to reach high fight intensities under flow conditions. It was possible to visualize O. rusticus urine release, and they released urine more often when upstream of an opponent in a flow environment during these agonistic bouts. These results suggest that O. rusticus collected from lotic environments release urine to maximize the transmission of chemical cues to a fight opponent. It appears that crayfish may adapt their signalling processes based upon their long-term ambient environments.     

The Effects of Starvation on Crayfish Responses to Alarm Odor

The effect of starvation on responses to alarm odor was tested with individuals of an invasive and a native species of crayfish. I predicted that chemical predator cues would inhibit feeding less in starving than well-fed animals, and that this decrease would be stronger in a native compared with an invasive species. Individuals were exposed to food odor and then alarm odor after 3 and 10 d of starvation. The inhibition of food-related behavior patterns was similar on the 2 d of testing for individuals of the invasive species, Orconectes rusticus. Individuals of O. virilis showed a significant reduction in the effects of alarm odor detection on day 10 compared with day 3 of starvation. The lack of a change in responses to alarm odors between days of testing by individuals of O. rusticus may be because they are more responsive to alarm odors than individuals of O. virilis. This behavioral difference could contribute to the successful range extension of O. rusticus.     

Complex predator-prey interactions and predator intimidation among crayfish,piscivorous fish,and small benthic fish

Summary Predator-prey interactions were studied among a small prey fish (the johnny darter Etheostoma nigrum) and two predators (crayfish Orconectes rusticus and smallmouth bass Micropterus dolomieui) with complementary foraging behaviors. When only smallmouth bass were present, darters reduced activity to 6% of control rates and spent most of the time hiding under tile shelters. When only crayfish were present, darter activity and shelter-use were similar to controls. When both crayfish and bass were present, an interaction occurred. Darters, normally inactive in the presence of bass, were often forced to move by approaching crayfish and thus activity increased to 19% of control rates. Also, darters were often evicted from shelters by intruding crayfish. Thus, crayfish increased the vulnerability of small fish to bass by evicting them from shelters and causing increased activity. Conversely, bass increased the vulnerability of small fish to crayfish by forcing these fish to seek cover under shelters occupied by crayfish. Intimidation effects of bass on darters last for some time. After a 30-min exposure to bass, darters showed reduced activity and increased shelter use lasting at least 24 h after the bass was removed. Thus predators, throught intimidation, can influence prey behavior even though the predators are no longer present.The Unit is sponsored jointly by the United States Fish and Wildlife Service, the Ohio Department of Natural Resources, and The Ohio State University     

Interactions among invaders: community and ecosystem effects of multiple invasive species in an experimental aquatic system

With ecosystems increasingly supporting multiple invasive species, interactions among invaders could magnify or ameliorate the undesired consequences for native communities and ecosystems. We evaluated the individual and combined effects of rusty crayfish (Orconectes rusticus) and Chinese mystery snails [Bellamya (=Cipangopaludina) chinensis] on native snail communities (Physa, Helisoma and Lymnaea sp.) and ecosystem attributes (algal chlorophyll a and nutrient concentrations). Both invaders are widespread in the USA and commonly co-occur within northern temperate lakes, underscoring the importance of understanding their singular and joint effects. An outdoor mesocosm experiment revealed that while the two invaders had only weakly negative effects upon one another, both negatively affected the abundance and biomass of native snails, and their combined presence drove one native species to extinction and reduced a second by >95%. Owing to its larger size and thicker shell, adult Bellamya were protected from crayfish attack relative to native species (especially Physa and Lymnaea), suggesting the co-occurrence of these invaders in nature could have elevated consequences for native communities. The per capita impacts of Orconectes (a snail predator) on native snails were substantially greater than those of Bellamya (a snail competitor). Crayfish predation also had a cascading effect by reducing native snail biomass, leading to increased periphyton growth. Bellamya, in contrast, reduced periphyton biomass, likely causing a reduction in growth by native lymnaeid snails. Bellamya also increased water column N:P ratio, possibly because of a low P excretion rate relative to native snail species. Together, these findings highlight the importance of understanding interactions among invasive species, which can have significant community- and ecosystem-level effects.     

Nest defense and aggressive interactions between a small benthic fish (the johnny darter Etheostoma nigrum) and crayfish

Synopsis Aggression by nest-guarding male johnny darters, Etheostoma nigrum, against intruding crayfish was investigated in laboratoy experiments and field observations. In the laboratory, darter success in chasing crayfish, Orconectes rusticus, from the nest site was inversely related to crayfish size. Small crayfish (less than 15 mm carapace length) were routinely evicted from the nest area by nips directed at the posterior end of the abdomen. Although such aggressive behavior was less successful against larger crayfish, even the largest crayfish tested (carapace length 30–32 mm) were chased from the nest area in 33% of the trials. Those large crayfish that entered nests often remained despite repeated attacks by the male johnny darter and egg predation was observed. In a small Ohio stream, openings to johnny darter nests were generally between 7 and 13 mm. Thus crayfish with a carapace height greater than 13 mm (corresponding to a carapace length greater than 29 mm) would not be able to enter johnny darter nests. In field observations, male Johnny darters successfully defended nests against another crayfish species (Orconectes sanborni, carapace length 12–29 mm). Together, aggressive behavior and small size of nest entrances allow the johnny darter to successfully reproduce in areas with abundant crayfish. The Unit is sponsored jointly by the United States Fish and Wildlife Service, The Ohio Department of NaturalResources, and The Ohio State University.