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.     

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.     

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.     

Biology and ecology of Thallomys nigricauda (Rodentia, Muridae) in the Thornveld savannah of South Africa

The black-tailed tree rat, Thallomys nigricauda, is one of the few arboreal rodents inhabiting the southern African savannahs. During three field campaigns in 2001 and 2002 we collected data on the biology and ecology of T. nigricauda in the Thornveld savannah of the Molopo Nature Reserve (NW Province, South Africa). T. nigricauda lives a solitarily lifestyle in the Molopo Nature Reserve. Diet analyses shows that they are folivorous, specializing on leaves of Acacia mellifera and Acacia luederitzii. Male and female animals have a similar body size (≈80 g). Mean age at weaning was 30 days and mean litter size at this age was 1.8. Monthly survival rates varied between 0.25 and 0.86. Density was estimated to be 0.5–1 individual ha⁻¹. Overall, our results show that the basic ecological characteristics of this arboreal rodent are fairly similar to other rodent species.     

Temporal shifts in physical habitat of the crayfish,Orconectes neglectus (Faxon)

A total of 300 samples was collected from February 1985 to August 1986 in a medium order Ozark Mountain stream. Physical habitat measurements of temperature, mean water column velocity, depth, and substrate character were recorded for each of the 25 monthly samples along with length and sex of all individuals of Orconectes neglectus (Faxon). Analysis of habitat utilization and suitability (or preference) was conducted using exponential polynomial models of hydraulic stress models. There appeared to be equal preference for depth over the range measured. Both substrate and velocity preference curves were bimodal with each mode designating certain crayfish size classes. Young-of-the-year were found primarily in cobbled, high velocity areas while adults were found in low velocity, macrophyte beds. Utilization curves for laminar sublayer thichness also reflected size-dependent phenomena where young-of-the-year were found in thin sublayer areas and adults were found primarily in thick sublayers. When separated by time and size, adults were found to occupy higher velocity, cobbled habitats during at least two months. This time period corresponded with the time of egg-bearing and further analysis yielded a time-dependent habitat suitability surface which accounted for this movement pattern. We suggest that the application of these suitability surfaces, which reflect habitat changes during the annual life cycle, will produce more accurate predictions of density and will allow better habitat management decisions under various regulated flow scenarios.     

A phasic-tonic proprioceptor in the telson of the crayfish Procambarus clarki (Girard)

Summary A proprioceptor in the telson of the crayfish Procambarus clarki was found, which is made up of about five units. These show range fractionation, i.e. different threshold distributions and their response to bending of the telson has a transient and a stationary component. The peak frequencies reached during the period of the transient component were shown to increase with increasing bending velocity and had — with the exception of one case — the highest values when extension (contrary to flexion) was imposed. The position characteristic discharge frequencies of the stationary component and their variation with different telson positions were considerably smaller. The receptor signals movement and position to the central nervous system and is therefore called a phasictonic one.I am grateful to Prof. Dr. T. H. Bullock for his hospitality and interest in this work and also want to thank Mr. John W. Thorson for many constructive discussions. This work was aided by grants from the U.S. Public Health Service, National Science Foundation and Office of Naval Research to Prof. Dr. T. H. Bullock.     

Effect of Temperature on Development and Immature Survival in the Scale Insect Predator, Chilocorus nigritus (F.) (Coleoptera: Coccinellidae)

Developmental and immature survival rates of the coccidophagous coccinellid Chilocorus nigritus (F.) were examined under constant, cycling and glasshouse temperatures in order to determine its suitability for use as a biological control agent in temperature glasshouses. First instar larvae did not complete development at 18 C. However, within the range 20-30 C, the developmental rate increased with rising temperature. The theoretical lower thermal threshold for development was found to be 16.6 C. Thermal summation and polynomial regression methods were used to predict developmental periods under glasshouse conditions. The predictions were accurate to within a mean of 10% in relation to observed data. Under laboratory conditions, immature survival rates were highest 28 C (52%) and lowest at (and below) 20 C (17%). First instar larvae suffered the highest mortality rates, while pupae had the lowest. Under glasshouse conditions, the survival rates were much lower (9% in the winter months and 20% throughout the remainder of the year), but the species was considered to be a suitable biocontrol agent if mean daily temperatures were maintained at levels above 20 C.     

A model for the time–temperature–mortality relationship in the chill-susceptible beetle, Alphitobius diaperinus, exposed to fluctuating thermal regimes

Exposing insects to a fluctuating thermal regime (FTR) compared with constant low temperature (CLT) significantly reduces cold-induced mortality. The beneficial effects of FTR result from physiological repair during warming intervals. The duration and the temperature experienced during the recovery period are supposed to strongly impact the resulting cold survival; however, disentangling the effects of both recovery variables had not been broadly investigated. In this study, we investigate cold tolerance (lethal time, Lt₅₀) of the polyphagous beetle Alphitobius diaperinus. We examined adult survival under various CLTs (0, 5, 10 and 15 °C), and under 20 different FTR conditions, where the 0 °C exposure alternated with various recovery temperatures (Rt) (5, 10, 15 and 20 °C) combined with various recovery durations (Rds) (0.5, 1, 2, 3 and 4 h). Under CLTs, Lt₅₀ increased with temperature until no mortality occurred above the upper limit of cold injury zone (ULCIZ). Under FTRs, Lt₅₀ increased with both Rt and Rd. The magnitude of the survival gain was clearly boosted when Rt was above the ULCIZ (at 20 °C). Based on a data matrix of lethal times with multiple Rt×Rd combinations, a predictive model showed that cold survival increased exponentially with Rt and Rd. This model was subsequently validated with additional survival tests. We suggest that increasing recovery durations associated with optimal recovery temperatures eventually leads to a progressive chilling compensation.     

The Behavioural Response of Mayfly Nymphs (Stenonema sp.) to Chemical Cues from Crayfish (Orconectes rusticus)

Artificial reef systems play an important role in the increase of natural production of biological marine resources and they have been deployed worldwide. In Portugal, seven artificial systems have been deployed along the southern coast of the Algarve. Research to date has focussed mainly on fish populations, particularly those of economical importance. The present work aims to study the macrobenthic communities of the artificial reef structures, as these will enhance the food resources and shelter, making the reefs more attractive to fish. In particular, we experimentally analysed the sequence of colonisation of macrobenthic communities of the Ancão artificial reef system, which was deployed in the summer of 2002. The study of the colonisation of benthic communities was done using samples of concrete cubic units (15 × 15 cm) that were suspended at the reef modules at a depth of 20 m, at the time of the reef immersion. Four replicate samples were collected by SCUBA diving from two groups of the Ancão reef every three months from the starting date. Sampling was done using essentially non-destructive methods to assess the percentage cover of macrobenthic organisms in both vertical and horizontal surfaces. The percentage cover of the taxonomic groups was compared within the different surfaces of the samples and between the two reef groups. The bottom surface of cubic samples had a significantly higher colonisation related to the dominance of barnacle cover, probably due to lower sedimentation levels. Samples from both reef groups showed a similar pattern of colonisation. Barnacles, bryozoans and serpulids dominated the samples three months immediately after the beginning of the experiment. Other invertebrates groups, such as Porifera, Hydrozoa, Anthozoa, other sessile Polychaeta, Decapoda, Gastropoda and Bivalvia, were more abundant after six months of colonisation.     

Directional selectivities of near-field filiform hair mechanoreceptors on the crayfish tailfan (Crustacea: Decapoda)

Directional selectivities of mechanoreceptors that innervate filiform hairs on the crayfish tailfan were investigated with unidirectional, sinusoidal, water-motion stimuli. These recordings provide the first representative sample from filiform hair sensilla on the entire tailfan. The filiform hair receptors exhibit unimodal directional selectivity patterns that were well fitted by a cardioid function with a half-width of 122°. The preferred directions correspond to the major axis of hair motion, and are perpendicular to the orientation of lateral branches on the main hair shaft. Pooled plots of preferred directions demonstrate quadrimodal patterns on the telson and endopods which are associated with hair location, and a bimodal pattern on the exopods. For each appendage, the combination of the overall pattern of preferred directions with “coarse coding” of direction by individual receptors provides sensitivity to a full 0–360° range of water motion and the potential to discriminate the direction of water motion throughout this range. The results demonstrate several similarities to the wind-sensitive cercal receptor system in orthopteroid insects, and suggest that crustacean filiform hair receptors provide a sufficient sensory basis for behavioral orientation to water currents and shorelines. Accepted: 5 January 1998     

Thermoregulation of spotted turtles (Clemmys guttata) in a beaver-flooded bog in southern Ontario,Canada

Body temperature has a major influence on the physiological processes, growth, reproductive output, and overall survival of ectotherms. When a habitat is altered as a result of natural or anthropogenic influences, the available temperatures in the habitat can change, thus affecting an animal's ability to thermoregulate. We studied thermoregulation in response to habitat change in a population of spotted turtles (Clemmys guttata) in Southern Ontario, Canada. Historically, the study site was ditched to draw down water levels to facilitate peat mining, and the resulting drainage ditches were the only habitat containing surface water and turtles were restricted to these drains. Recent colonization of the site by beaver (Castor canadensis) caused increases in water level and water surface area. We followed spotted turtles (N=16) outfitted with radio transmitters and iButtons to estimate body temperatures (T_b) continuously throughout the active season post-flooding. Turtle models outfitted with iButtons (N=50) were deployed in the nine available habitat types to record environmental temperatures (T_e). Turtles (N=13) were tested in a thermal gradient under laboratory conditions to determine preferred body temperature range (T_set). The T_set for the population ranged from 20 °C to 26 °C. In the field, T_b was within the T_set range 28% of the time from March to October, and 67% of the time from July to August. Efficiency of thermoregulation was calculated to be highest in July and August. The habitat type with the highest thermal quality was the shallow flooded zone created by beaver damming, and the habitat with the lowest thermal quality was the drain bottom, the drains being the only aquatic habitat available prior to flooding. This study confirms that beaver flooding provided a wide variety of preferable thermal opportunities for spotted turtles. Further investigation is needed to determine the effects of flooding on spotted turtle thermoregulation during nesting and hibernation.     

Evolution of thermal physiology and growth rate between populations of the western fence lizard (Sceloporus occidentalis)

Summary Hatchling Sceloporus occidentalis from northern populations (central Oregon) grow more slowly than hatchlings from southern populations (southern California) in nature. In this study, I determine whether this difference in growth rate results from differences in thermal environment and/or in thermoregulatory behavior. To determine the degree to which the thermal environment affects growth rate among populations, I reared hatchings from the northern and southern populations in a cycling thermal regime in one of three experimental treatments differing in access to radiant heat (6, 9, or 12 h radiant heat; remainder of 24 h at 15°C). I also measured the body temperature that each individual voluntarily selected over the course of the daily activity cycle. Growth rate varied positively with duration of access to radiant heat. Within the three treatments, individual growth rate was positively correlated with body temperature. Moreover, the difference in growth rate between the northern and southern populations was due in part to differences in behavior — individuals from northern populations selected lower body temperatures. I found that significant variation in body temperature was associated with family membership, suggesting that thermal physiology has a genetic basis. Moreover, growth rate was correlated with body temperature among families in each population suggesting a genetic correlation underlies the phenotypic correlations. Thus, genetically based variation in thermal physiology contributes to differences in growth rate among individuals within a population as well as to differences among populations.     

Effects of cold- and heat hardening on thermal resistance in Drosophila melanogaster

The effects of cold- and heat hardening on resistance to both low and high temperature stress was examined in Drosophila melanogaster lines selected for resistance to either cold or heat. The hardening effect was positive when the hardening was of the same type as the stress in all selection regimes. The effect of cold hardening on survival after heat stress was further examined in the lines selected for cold resistance and corresponding controls. A cross-protection effect (increased heat resistance after cold hardening) was present and this effect was lower in the lines selected for resistance to cold than in the controls. The level of Hsp70 expression induced by a non-lethal cold hardening was examined, showing that cold hardening induced Hsp70 expression. The results suggest that the cross-protection effect is at least partly due to Hsp70 expression induced by cold exposure.     

Fighting Strategies in Crayfish Orconectes rusticus (Decapoda, Cambaridae) Differ with Hunger State and the Presence of Food Cues

Crayfish, bearing dangerous weapons in the form of chelae, resolve intraspecific conflicts using stereotyped behaviors and structured, escalated encounters. According to predictions of game theory models, any decision to resort to unrestrained combat without prior careful behavioral assessment of the opponent's fighting abilities carries great risks. The present study examines the significance of internal hunger states and the presence of chemical food cues in this decision process using a 2 × 2 factorial design. Hungry crayfish escalated more rapidly, and thus took greater risks, during agonistic encounters, while the presence of a food source reduced the rate at which fights increased in intensity. However, there were no significant differences in fighting behavior as a result of the interaction between these two variables. We then address the complex trade-offs that individuals face in fighting with respect to increased risks of injury, appetitive states, and opportunities for resource access.     

Mortality curves of blind cave crayfish (Orconectes australis australis) exposed to chlorinated stream water

Chlorinated stream water toxicity was tested on the blind cave crayfish Orconectes australis australis from Merrybranch Cave, White County, Tennessee. An undisturbed natural cavern, Merry-branch was formed between two strata of sandstone having a mean elevation of 354 meters (MSL). Test water was collected from a subterranian stream in the cave supporting the hypogean crayfish population, and transported to the laboratory. No chlorinity was detected in the underground stream water.In the laboratory, cave water was chlorinated with sodium hypechlorus solution at various concentrations of total residual chlorine, combined residual, and free chlorine content as measured by Standard Methods titration procedure. Thirty-six crayfish, six crayfish per test solution, were subjected to a three day acclimation period at chlorine concentrations ranging from 0.21–1.50 mg./l. total residual chlorine, 0.20–0.30 mg./l. combined residual chlorine, and 0.01–1.20 mg./l. free chlorine; and then subjected t0 a 24 hour time-to-death (hourly) bioassay at the following chlorine water dilutions (mg./l.): (1) 7.45 total residual, 0.45 combined residual, and 7.00 free, (2) 3.39 total residual, 0.39 combined residual, and 3.00 free, (3) 2.85 total residual, 0.35 combined residual, and 2.50 free, (4) 2.30 total residual, 0.30 combined residual, and 2.00 free, (5) t.96 total residual, 0.21 combined residual, and 1.75 free, and (6) control. Fluctuations within these concentrations ranged from ± 0.20 free chlorine. All test solutions and a control were delivered by an Esvelt serial diluter. In addition, a 24 hour time-to-death (hourly) bioassay was conducted at the same dilutions 0n crayfish not acclimated to chlorine.These test demonstrated that crayfish mortalities generally increased with increasing concentrations of chlorine in both bio-assays, while acclimated crayfish tended to be more tolerant than non-acclimated ones.Funds for this research were provided by the Aquatic Ecology Fund, Biology Dept., Tennessee Technological University     

Cross-species amplification of microsatellite markers in the invasive spiny-cheek crayfish (Orconectes limosus): Assessment and application

The North American spiny-cheek crayfish,Orconectes limosus (Cambaridae), endangered in its native range, is a widespread invasive species in European waters and conservationally important carrier of crayfish plague. However, its population structure is poorly known, and no informative genetic markers for the species are available. We tested cross-species transfer of microsatellite loci to spiny-cheek crayfish from 5 other crayfish species. Variability of 10 successfully amplifying loci derived from 4 species was then tested in 60 individuals ofO. limosus originating from 3 natural populations: the river Danube at Bogyiszló in Hungary, a pond in Stary Klíčov, and the brook Černovicky, both in the Czech Republic. The allele number within the populations ranged from 4 to 10 alleles per locus, while heterozygosity levels varied from 0.650 to 0.900 forHo and from 0.660 to 0.890 forHe. No linkage disequilibrium and no null alleles were detected. The selected markers are useful for assessing population structure, intraspecific variation, and paternity studies in spiny-cheek crayfish.     

Isolation of the rhabdomeral microvillar cytoskeleton of the crayfish (Orconectes limosus) photoreceptor by a crosslinking reagent.

1. Rhabdomeral microvilli of photoreceptor cells of invertebrates contain a labile central cytoskeleton. For stabilization of the rhabdomeral cytoskeleton of the crayfish Orconectes limosus the crosslinking reagent suberic acid bis (N-hydroxysuccinimide ester) was used. 2. It was found that this crosslinking reagent can be successfully used to stabilize and isolate the microvillar cytoskeleton of crayfish photoreceptors. 3. After detergent treatment cytosolic proteins and the cell membranes were removed. 4. By the combined use of crosslinker and detergent the accessibility of antibodies or other markers to the microvillar cytoskeleton is possible. 5. This method may be useful, because at present little is known about the proteins associated with the central filament of invertebrate photoreceptors.     

Size-biased predation by the gila topminnow poeciliopsis occidentalis (Baird and Girard)

The objective of this study is to document under both field and laboratory conditions the phenomenon of size-biased feeding by the Gila topminnow, Poeciliopsis occidentalis, and to shed light on the mechanisms responsible for this process. Adult female Gila topminnows feed in Monkey Spring, Santa Cruz County, Arizona, on a variety of food items, but Hyalella azteca is an important element of its invertebrate diet. The size of the amphipods ingested by female topminnows is a function of the size of the fish, with larger fish taking larger prey. There is also a strong preference for topminnows to select large amphipods from within the size limits imposed by the width of the mouth.The correlation between fish size and prey size was also seen in the laboratory, utilizing Daphnia similis and food pellets as prey. The tendency to select large prey from a range of available sizes was prominent with both Daphnia and pellets. The feeding period in the laboratory is characterized by two distinct phases. During the first phase, feeding is rapid and nonselective. As the feeding progresses, the rate of feeding drops dramatically and selectivity increases.More complex laboratory feeding experiments gave equivocal results. When fish were presented pellets of two sizes at different densities and at different ratios of small to large, size biased feeding was not clearly evident. Both predator and prey densities, agonistic behavior, and reaction distance should be explored more thoroughly as next steps in attempts to mimic feeding behavior in nature.Extracted from a Ph. D. thesis written by the junior author     

Effect of temperature and thermal acclimation on locomotor performance of Macrobiotus harmsworthi Murray (Tardigrada, Macrobiotidae)

We investigated the effects of acute and acclimation temperature on the locomotor performance and behavior of the tardigrade Macrobiotus harmsworthi collected from Qinling Mountains in central China. Tardigrades were acclimated to either 10 or 25 °C for 2 weeks. Then we recorded their walking speed, percentage of time moving, and the maximum distance covered by continuous locomotion at either 10 or 25 °C as the rate parameters of locomotor performance. The walking speeds of M. harmsworthi varied from 1.98 to 4.8 mm min^–1. The locomotor performance rates were significantly influenced by both acclimation temperature and performance temperature and by the interaction of the performance temperature and acclimation temperature. The data from our studies support the Beneficial Acclimation Hypothesis (BAH) which predicts that animals acclimated to a particular temperature have enhanced performance or fitness at that temperature in comparison with animals acclimated to other temperatures. The data, at least potentially, also support the Warmer is Better Hypothesis which predicts that organisms raised at high temperatures have higher relative fitness across all temperatures than do those raised at intermediate or cool temperatures. Some of the results from our studies testify the inference from the BAH that performance temperature that deviates from the acclimation temperature could cause the reduction of the locomotor performance rate.