Predator Recognition and Learned Irrelevance in the Crayfish Orconectes virilis

When individuals of the crayfish Orconectes virilis detect an unlearned danger cue (alarm odor) and a novel cue (goldfish odor) at the same time, they form a learned association and behave as if the novel cue is associated with increased predation risk ( Hazlett et al. 2002 ). This study examined the potential for learned irrelevance in O. virilis and the circumstances under which blockage of the formation of a learned association could occur. If individuals experience a random pattern of alarm odor and goldfish odor over the days prior to the simultaneous detection of those two cues, no learned association is formed (= learned irrelevance). That is, there is no inhibition of responses to a food cue when goldfish odor is added if the crayfish has experienced a random pattern of the two cues. Learning was eliminated if the random pattern of cues was experienced before or after the simultaneous detection. To present the two cues (alarm and goldfish odors) to crayfish independently on separate days, the water containing goldfish odor had to be removed from the aquaria as the odor persisted at least 24 h. The importance of the learned irrelevance phenomenon on predator–prey interactions is discussed.     

The impact of the crayfish Orconectes virilis on aquatic macrophytes

SUMMARY. 1. The impact of crayfish on the biomass, density and shoot morphology of four submersed plant species was examined under semi-natural conditions. Male or female crayfish ( Orconectes virilis ) were held for 5 weeks at biomasses of 0, 5, 10 or 18 g m⁻² (live weight) in twelve plastic pools (4.67 m², surface area) containing Potamogeton richardsonii, Myriophyllum exalbescens, Nuphar variegatum and Sparganium eurycarpum .
2. Crayfish significantly affected biomass, density and/or shoot morphology of all four macrophyte species. Differences in the effect of crayfish on macrophyte growth were related to plant species, crayfish sex and activity, and the abundance of alternative foods.
3. The effect of female crayfish on macrophyte growth was generally stimulatory. Myriophyllum and Potamogeton biomass, Potamogeton density and Myriophyllum length increased in the presence of female crayfish, possibly due to the reduction in herbivorous snails as a result of crayfish predation. In contrast, plant growth decreased in the presence of male crayfish: Myriophyllum, Nuphar and Potamogeton biomass, Myriophyllum and Sparganium density, and Sparganium and Poiamogeton length were reduced at male crayfish biomasses between 5 and 18 g m⁻².
4. These results indicate that even relatively low densities of crayfish can greatly affect the growth of submersed aquatic plants. Because of their ability to modify aquatic macrophyte, macroinvertebrate and, ultimately, fish communities, the introduction of crayfish into lakes where they do not occur could have a major effect on the structure and composition of the littoral zone.     

The Influence of Dominance on Shelter Preference and Eviction Rates in the Crayfish, Orconectes rusticus

The establishment and reinforcement of dominance status is thought to provide an individual with increased access to preferred resources, such as food, mates, and shelter. Resources within an environment vary based on their availability, abundance, and specific characteristics of the resource. Animals can exhibit preferences for resources by increasing the intensity of competition for access to the resource. We investigated the role that dominance has in acquiring and controlling resources within crayfish populations. Populations of 5 size‐matched crayfish were allowed to establish a social hierarchy in the presence of shelters of differing structure in 10 different trials. Crayfish agonistic interactions and the use and control of shelters were quantified from 96 h continuous video recordings. Shelter preference was defined by crayfish use of specific shelter types that were occupied for longer durations. Analysis of the social hierarchy demonstrated that crayfish have a linearity hierarchy in this context. In addition, it is clear that dominance rank had a significant impact on shelter use, shelter acquisition, and shelter eviction. Crayfish with lower average dominance ranks selected specific shelter types (smaller and fewer openings). Dominant crayfish used big shelters and were less likely to be evicted from shelters. These results demonstrate that dominance in crayfish serves a role in the acquisition of resources and is also important for the control of those resources.     

Is Competition with the Invasive Crayfish Orconectes neglectus chaenodactylus Responsible for the Displacement of the Native Crayfish Orconectes eupunctus?

The crayfish Orconectes eupunctus, a species of special concern, appears to have been displaced from part of its former range in the Spring River drainage of Arkansas and Missouri by the invasive crayfish O. neglectus chaenodactylus. We conducted preliminary field experiments to determine whether Orconectes eupunctus were displaced from their former range and whether interspecific competition with O. neglectus chaenodactylus was responsible for the displacement. To determine whether Orconectes eupunctus were displaced from their former range by negative biotic interactions with other species we placed single adult male O. eupunctus in cages at two sites: an upstream site where O. eupunctus was formerly abundant and a downstream site where it is currently abundant. Orconectes neglectus chaenodactylus were additionally paired alongside O. eupunctus at the upstream site. We also conducted a field competition experiment with treatments consisting of (1) 3 O. eupunctus, (2) 6 O. eupunctus, and (3) 3 O. eupunctus and 3 O. neglectus chaenodactylus at a site where both species occur. In the first experiment, O. eupunctus gained significantly more weight at the downstream site than the upstream site during 2003, but not during 2002. At the upstream site, O. eupunctus showed significantly greater growth rates than O. neglectus chaenodactylus during 2002, but not during 2003. In the competition experiment, O. neglectus chaenodactylus did not significantly reduce the growth or survival of O. eupunctus. These results suggest that O. eupunctus have been displaced from their former range by biotic interactions, but interspecific competition between adult male O. eupunctus and O. neglectus chaenodactylus does not appear to be the mechanism responsible for this displacement. Further study is needed to investigate the disappearance of O. eupunctus.     

The tradeoff between reproductive and food resources in the crayfish Orconectes virilis

The conflict between mating and foraging is one of the many tradeoffs associated with reproduction. The relative attraction to sex pheromone and food stimuli by male virile crayfish (Orconectes virilis) were tested using two experiments. In the first experiment, the responses by males to pheromone and food stimuli alone were assessed. In the second experiment, males were exposed to both pheromone and food stimuli in a choice maze. To determine the effects of feeding motivation, the males in the choice experiment were fed either one day or seven days prior to observation. Male crayfish exhibited similar responses to the pheromone and food stimuli individually. Males exhibited a preference for the pheromone stimulus over the food stimulus in the choice trials, and that preference that was not influenced by starvation.     

The mate pursuit-predation avoidance tradeoff in the virile crayfish, Orconectes virilis

To better understand the tradeoff between mate pursuit and predation risk avoidance, the responses of male virile crayfish (Orconectes virilis) to female sex pheromone and conspecific alarm cue, alone and in combination were tested. Male crayfish were more active and exhibited less antipredator behavior in the pheromone treatment than in either the pheromone + alarm or alarm treatment. The results suggest a tradeoff in favor of risk avoidance when pheromone and risk stimuli are encountered together.     

The mate pursuit-predation avoidance tradeoff in the virile crayfish,Orconectes virilis

To better understand the tradeoff between mate pursuit and predation risk avoidance, the responses of male virile crayfish (Orconectes virilis) to female sex pheromone and conspecific alarm cue, alone and in combination were tested. Male crayfish were more active and exhibited less antipredator behavior in the pheromone treatment than in either the pheromone?+?alarm or alarm treatment. The results suggest a tradeoff in favor of risk avoidance when pheromone and risk stimuli are encountered together.     

Dichroism of Photosensitive Pigment in Rhabdoms of the Crayfish Orconectes

Microspectrophotometric measurements of isolated crayfish rhabdoms illuminated transversely show that their photosensitive absorption exhibits a dichroic ratio of 2 in situ. The major absorption axis matches the axial direction of the closely parallel microvilli comprising the receptor organelle. Since these microvilli are regularly oriented transversely in about 24 layers, with the axes of the microvilli at 90° in alternate layers, transverse illumination of a properly oriented rhabdom displays alternate dichroic and isotropic bands. Because all the microvilli from any one cell share the same orientation, the layers of microvilli constitute two sets of orthogonal polarization analyzers when illuminated along the normal visual axis. Furthermore, since the dichroic ratio is 2 and transverse absorption in isotropic bands is the same as that in the minor absorbing axis of dichroic bands, the simplest explanation of the analyzer action is that the absorbing dipoles of the chromophores, as in rod and cone outer segments, lie parallel to the membrane surface but are otherwise randomly oriented. The rhabdom's functional dichroism thus arises from its specific fine structural geometry.     

Selective foraging by the crayfish Orconectes virilis and its impact on macroinvertebrates

SUMMARY. 1. The impact of crayfish predation on the abundance of macroinvertebrates was examined under semi-natural conditions. Female (Experiment 1) or male (Experiment 2) crayfish (Orconectes virilis) were held for 5 weeks in twelve small pools (4.67 m² surface area) at biomasses of 0. 5, 10 or 18 g m^?2 (live weight). The pools were stocked with known densities of macroinvertebrates. 2. Crayfish significantly affected the abundance of macroinvertebrates in the pools. Differences in the effects of crayfish on macroinvertebrates were related to crayfish sex, the presence of age-0 crayfish, and the species of macroinvertebrate. 3. The abundance of snails (Stagnicola elodes and Physa gyrina) was greatly reduced, in comparison with controls, by biomass of female crayfish ≥10 g m^?2 and by biomasses of male crayfish ≥5 g m^?2. The total density of non-molluscan invertebrates was inversely correlated with the biomass of female crayfish but the total biomass of non-molluscan invertebrates did not differ between treatments. This is consistent with our observation that small invertebrates (<2 mg wet weight) were less numerous, and large amphipods (32–64 mg) were more numerous, in pools stocked with female crayfish. In contrast, male crayfish had little apparent effect on the abundance of non-molluscan invertebrates. 4. Age-0 crayfish hatched at the end of Experiment 1 and were present in each pool at the start of Experiment 2. Surprisingly, male crayfish preyed little on age-0 crayfish. At the end of Experiment 2, the densities of age-0 crayfish varied between six and 116 individuals m^?2 and there was a strong inverse correlation between the mean biomass and density of age-0 crayfish recovered from the pools. This suggests age-0 crayfish were food limited in the pools and may explain the dominance of oligochaetes (which largely escape predation by burrowing) in the invertebrate community at the end of Experiment 2. 5. These results indicate that even relatively low densities of crayfish could greatly affect the abundance of macroinvertebrates in lakes. The introduction of crayfish into lakes (most lakes in Alberta currently have no crayfish) could substantially affect abundance and species composition of the macroinvertebrate community and, ultimately, the fish populations.     

Some factors regulating cohort production of the crayfish, Orconectes virilis

SUMMARY. Cohort production of the crayfish. Orconectes virilis , ranged from 148.8 to 70.2 kg in North Twin Lake (surface area 1.9 ha) and from 151.9 to 84.9 kg in West Lost Lake (surface area 1.4 ha). Both cohort production (P) and cohort mean biomass (B) showed a long-term decrease with time. The cohort P/B ratios did not show such a trend but remained steady. The P/B ratio approximated the life cycle instantaneous rate of growth (G) which could then be used for estimating cohort production from mean biomass.
Production remained consistently higher in West Lost Lake. Analysis of horizontal life table data revealed no inherent difference between survival and fecundity rates of the crayfish population in each lake. Slight differences in basin morphometry between the two lakes determined nursery habitat availability and this affected recruitment of young from each female brood stock. Small differences in carrying capacity between two similar sized nearby lakes, along with a decrease in carrying capacity over time, were sufficient to nullify attempts to manage these populations using a maximum sustainable yield model.     

Intercellular junctions of antennal gland epithelial cells in the crayfish,Orconectes virilis

Summary Labyrinth and nephridial canal cells of the crayfish (Orconectes virilis) antennal gland possess two types of intercellular junctions revealed by freeze-fracture studies. Apical margins of the cells are connected by long septate junctions. In replicas, these junctions consist of many parallel rows of 80–140 Å intramembrane particles situated on the PF membrane face (EF and PF fracture faces of Branton et al., 1975). Rows of pits are found on the EF fracture face and are deemed complementary to the rows of particles. Moreover, lateral margins of basal regions of the epithelial cells are attached by many intercellular junctions. These contacts are characterized in thin plastic sections by a narrow dense cytoplasmic plaque located subjacent to the plasma membrane at sites of adjoined cells, and 5 to 12 fine strands of dense material that extend across the intercellular gap between adjoined cells. In freeze-fracture replicas, EF intramembrane faces basal to the region of the plasma membrane containing septate junctions exhibit numerous discoid clusters of particles. The particle aggregates, assumed to represent freeze-cleave images of adhering junctions, range from 900 to 3,700 Å in diameter, with individual particles about 185 Å in diameter. These junctions appear to connect epithelial cell processes formed by basal infoldings of the plasma-lemma, and occur between adjacent cells as well as adjacent processes of a single cell. The discrete aggregates of particles resemble replicated desmosomes (Shienvold and Kelly, 1974) and hemi-desmosomes (Shivers, 1976); therefore, they probably do not constitute a basis for electrical coupling between antennal gland epithelial cells.Supported by the National Research Council of Canada     

Cytochemical observations of hemolymph cells during coagulation in the crayfish, Orconectes virilis

Smears of Orconectes virilis hemolymph were prepared by fixation in 10% formalin immediately upon withdrawal, and after an interval of 1, 5, or 15 minutes. These smears were then stained by a variety of histochemical means designed to identify a material which is released from both hyaline and granular cells during the coagulation process. Based upon reactions to various carbohydrate, lipid and protein tests, this material appears to consist of a glycoor muco-protein, and its localization and activity coincide with the ?fibrin ferment”? described by Hardy (1892). Its release is probably initiated from the hyaline cells, followed shortly thereafter by a similar activity of the granulocytes. While it cannot be ascertained by the methods used in this study whether or not this material participates actively in clotting, its release is nevertheless coincidental with coagulation.     

Histological and histochemical observations of the hemolymph cells in the crayfish, Orconectes virilis

Hemolymph cells of Orconectes virilis were stained during the months of August to November by a variety of histological and histochemical techniques. Cells were classified as hyaline cells, small granulocytes, and large granulocytes. Presence of mitochondrial enzymes was indicated by tests for succinic dehydrogenase and cytochrome oxidase. Reaction to test for the hydrolytic enzyme, leucine acylnaphthylamidase was intense in the granules of the large granulocytes. The PAS reaction indicated a mucopolysaccharide at the cell membrane. Lipid was found in all three hemolymph cell types of Orconectes virilis at the time of this study.     

The effect of aquatic plant chemistry and morphology on feeding selectivity by the crayfish, Orconectes virilis

SUMMARY. 1. Feeding preference trials conducted with male and female crayfish ( Orconectes virilis ) showed that out of ten submersed aquatic plant species, Chara sp. and Lemna trisulca were the preferred food sources, followed by Myriophyllum exalbescens or Utricularia vulgaris .
2. Analysis of plant fibre, alkaloid, total phosphorus, total and available nitrogen, and organic content showed that crayfish feeding preferences were not related ( P >0.5) to plant fibre or alkaloid content but were negatively correlated ( P <0.05) with plant nutrient content such that species of low nutritional value ( Chara and Lemna ) were consistently preferred.
3. Despite this preference for low nutrient food, daily intake rates by crayfish for phosphorus, nitrogen and organic matter were greatest when feeding on Lemna or Chara due to the large quantity consumed.
4. We suggest that feeding preferences of O. virilis are not determined by plant chemistry but, rather, relate to ease of handling small, short, bottom-dwelling plants such as Chara and Lemna as compared with large erect, roseate or floating-leaved forms. Because crayfish are selective herbivores, their introduction into waters where they do not naturally occur could have a dramatic effect on both the species composition and biomass of submersed plants.     

Exploration of potential microbial control agents for the invasive crayfish,Orconectes virilis

The introduced crayfish, Orconectes virilis (Cambarinae; Hagen 1870), has become a serious invasive species in Arizona (USA), altering stream ecosystems and contributing to the decline of native species. But because it is native to the eastern US, and related crayfish including endangered species inhabit nearby states, introduction of a biological control agent presents a unique challenge. This represents a feasibility study, in which we explored bacteria, nematodes, and a virus as potential biological control agents for O. virilis while avoiding harm to native species. White Spot Syndrome Virus (WSSV) from shrimp was found to be highly pathogenic and readily passed by cannibalistic behavior but not by water transmission. Potential activity of WSSV against representative non-target arthropods that may be found in the same ecosystems was also explored.     

Changes in Odour Intermittency Influence the Success and Search Behaviour During Orientation in the Crayfish (Orconectes Rusticus)

Aquatic organisms may use different mechanisms to locate an odour source, including odour-gated rheotaxis and chemotaxis. However, the effects of spatial and temporal information contained within odour signals in guiding orientation are under debate. In nature, turbulence and plume meandering give rise to signal intermittency, which organisms must overcome to locate an odour source successfully. In this study, crayfish were presented with odour plumes formed by a continuously released jet or a pulsed jet ranging from 0.5 to 3 Hz. Crayfish oriented significantly differently in pulsed odour plumes differed significantly from than in a continuously released jet plume. In a continuous odour plumes, crayfish walked faster, had more accurate heading and turning angles, and stopped less often. The results suggest that crayfish are able to orient in a pulsed odour plumes, but that the spatial and temporal complexity of a continuous released odour plume allowed them to be more successful to find the odour source.     

Field observations of intraspecific agonistic behavior of two crayfish species,Orconectes rusticus and Orconectes virilis,in different habitats

Agonistic behavior is a fundamental aspect of ecological theories on resource acquisition and sexual selection. Crustaceans are exemplary models for agonistic behavior within the laboratory, but agonistic behavior in natural habitats is often neglected. Laboratory studies do not achieve the same ecological realism as field studies. In an attempt to connect laboratory results to field data and investigate how habitat structure affects agonistic interactions, the nocturnal behavior of two crayfish species was observed by scuba diving and snorkeling in two northern Michigan lakes. Intraspecific agonistic interactions were analyzed in three habitats: two food resources-macrophytes and detritus-and one sheltered habitat. The overall observations reinforce the concept that resources influence agonistic bouts. Fights in the presence of shelters were longer and more intense, suggesting that shelters have a higher perceived value than food resources. Fights in the presence of detritus patches had higher average intensities and ended with more tailflips away from an opponent, suggesting that detritus was a more valuable food resource than macrophytes. In addition, observations of aggressive behavior within a natural setting can add validity to laboratory studies. When fights in nature are compared with laboratory fights, those in nature are shorter, less intense, and less likely to end with a tailflip, but do show the fundamental fight dynamics associated with laboratory studies. Extrinsic and intrinsic factors affect intraspecific aggression in many ways, and both should always be recognized as having the potential to alter agonistic behavior.     

Sexual History Affects Mating Behavior and Mate Choice in the Crayfish Orconectes limosus

Because mating entails both costs and potential benefits to both sexes, males and females should be under selection to make optimal choices from among available potential mates. For example, in some cases, individuals may benefit by using information on potential mates' previous sexual histories to make mate choices. In such cases, the form and direction of these benefits may vary both between the sexes and based on the sexual history of the choosing individuals themselves. We investigated the effects of recent previous sexual history on the mate choice and mating behavior of both males and females of the crayfish Orconectes limosus. In one experiment, we found that opposite‐sex dyads comprising crayfish that had both mated 7–8 d previously with other conspecifics were significantly less likely to mate than dyads in which at least one crayfish was unmated. In a second experiment, we found that, when presented with a choice of tethered (but free to move) opposite‐sex conspecifics, only virgin females discriminated between males based on sexual history, showing a preference for virgin males over recently mated males. Mated females, mated males, and virgin males showed no preferences based on the sexual histories of potential mates. We discuss the implications of these inferences in the context of what was previously known about mating behavior and potential sperm limitation in crustaceans and other taxa.