Freshwater crayfish invasions: former crayfish invader Galician crayfish hands title “invasive” over to new invader spiny-cheek crayfish

Biological invasions are a major threat to global biodiversity. Invasive freshwater crayfish in that context are especially prominent for their negative effects on both ecosystem integrity and native crayfish. However, some systems may have supported a crayfish species not originally native to the system without perceivable negative consequences for the ecosystem while other invasive crayfish species may constitute a major threat to ecosystem stability. Here I present an example how two crayfish, the spiny-cheek and the Galician crayfish both by researchers and governmental agencies considered non-native differ in their threats to the native ecosystem. Whereas the spiny-cheek crayfish is a recent potentially disease-transmitting and still spreading invader with high local densities the Galician crayfish might be part of the lake’s fauna since several hundred years, appears in lower densities and is unlikely to be a vector of disease. Therefore, regardless of the Galician crayfish’s actual date of introduction it is thus a rather “old and integrated” invader, which is now being faced and itself potentially threatened by the emergence of a “new and dangerous” invader: the spiny-cheek crayfish. This also exemplifies that in the face of often insufficient scientific information about dates of species introductions care should be taken in postulating species as invasive and dangerous without any form of risk assessment for their impact on the ecosystem.     

Does the introduced signal crayfish occupy an equivalent trophic niche to the lost native noble crayfish in boreal lakes?

The introduced North-American signal crayfish (Pacifastacus leniusculus) has become widespread throughout Europe where it has often replaced the native noble crayfish (Astacus astacus). The impact of this replacement on ecosystem processes in boreal lakes is still largely unknown. We compared the trophic niches of these two crayfish species in 16 small to medium sized boreal lakes in southern Finland; eight lakes with noble crayfish and eight lakes where the native crayfish populations had been lost and replaced by signal crayfish. We analysed carbon and nitrogen stable isotopes from samples of the crayfish and their putative food sources, and used stable isotope models to compare trophic niche widths of the two species of crayfish and to quantify the food sources used by them. At species level the signal crayfish exhibited a substantially larger trophic niche than that of the noble crayfish, but within-lake populations of the species did not differ in their niche widths. The isotopic niches of the two species strongly overlapped, and while the estimated proportions of food resources (profundal and littoral macroinvertebrates, terrestrial leaf detritus and macrophytes) used by crayfish varied considerably among individual populations, they did not differ consistently between the species. Our results suggest that, contrary to often expressed concerns, replacement of lost noble crayfish populations by the signal crayfish may not greatly alter the littoral food web structure in boreal lakes.     

Plasticity in fecundity highlights the females’ importance in the spiny-cheek crayfish invasion mechanism

Invasion is one of the most consequential phenomena affecting the distribution of native species. Few in number of species, European crayfish are losing the competition with introduced North American crayfish. The spiny-cheek crayfish, Orconectes limosus, is an outstanding example, successfully competing against the native narrow-clawed crayfish, Astacus leptodactylus. For four years, we collected data regarding crayfish occurrences, their relative abundance, and the structure of populations in the ongoing colonisation process of O. limosus in the lower Danube. The mature females of both invasive and indigenous crayfish species were analysed with respect to biometry and production of oocytes in relation to the dynamics of invasion. The interspecific comparisons showed no significant differences regarding body size, with an average of approximately 102 mm total length and 31 g wet weight for both species. However, the fecundity of the indigenous species was found to be constant throughout the investigated area, whereas the number of eggs produced by the invasive females was significantly increased at the active front of the invasion. The maximum number of ovarian eggs found was 887 and 1156 in the indigenous species and the invasive species, respectively. We propose the scenario that the invasive species, which carries the deadly crayfish plague, creates an ecological advantage by reducing the populations of indigenous crayfish. Subsequently, the invasive females opportunistically use the available resources to enhance their fecundity, resulting in the acute growth of populations. However, the long-term competitiveness and colonisation success of O. limosus still remain in question.     

Patterns in benthic food webs: a role for omnivorous crayfish?

1. The biomass and species richness of macrophytes and invertebrates in artificial ponds at two sites in southern Sweden (twenty-one ponds at each site) were investigated. Alkalinity was high at one site (H ponds) and low at the other site (L ponds). The ponds chosen had different densities of signal crayfish (Pacifastacus leniusculus), with mean crayfish abundance (estimated by trapping and expressed as catch per unit effort) significantly higher in the L ponds (10.7) than in the H ponds (4.9). Macrophytes, invertebrates, the amount of periphyton on stones and the organic content of the sediment were determined in each pond. 2. Macrophyte biomass, cover and species richness declined with increasing crayfish density. Macrophyte species composition differed between ponds and was related to crayfish abundance. 3. The total biomass of invertebrates and the biomass of herbivorous/detritivorous invertebrates declined with increasing crayfish abundance, but the biomass of predatory invertebrates declined only in the L ponds. The relative biomass of Gastropoda and Odonata declined in ponds where crayfish were abundant. In ponds where crayfish were abundant the invertebrate fauna was dominated by sediment-dwelling taxa (Sialis (H and L ponds) and Chironomidae (H ponds)). 4. The number of invertebrate taxa in macrophytes declined with increasing crayfish abundance. The percentage of macrophyte-associated invertebrate taxa differed between ponds, but also between sites. The relative biomass of Gastropoda declined in H ponds where crayfish were abundant. In H ponds Trichoptera or Gammarus sp. and Heteroptera dominated where crayfish were abundant, whereas Odonata dominated in L ponds with abundant crayfish. 5. The organic content of the sediment decreased in ponds with high crayfish densities, while the amount of periphyton on stones was not related to crayfish density. 6. We conclude that the signal crayfish may play an important role as a keystone consumer in pond ecosystems, but lower trophic levels did not respond to changes in the abundance of the crayfish according to the trophic cascade model. Omnivorous crayfish may decouple the cascading effect.     

Physiological thermoregulation in a crustacean? Heart rate hysteresis in the freshwater crayfish Cherax destructor

Differential heart rates during heating and cooling (heart rate hysteresis) are an important thermoregulatory mechanism in ectothermic reptiles. We speculate that heart rate hysteresis has evolved alongside vascularisation, and to determine whether this phenomenon occurs in a lineage with vascularised circulatory systems that is phylogenetically distant from reptiles, we measured the response of heart rate to convective heat transfer in the Australian freshwater crayfish, Cherax destructor. Heart rate during convective heating (from 20 to 30 degrees C) was significantly faster than during cooling for any given body temperature. Heart rate declined rapidly immediately following the removal of the heat source, despite only negligible losses in body temperature. This heart rate 'hysteresis' is similar to the pattern reported in many reptiles and, by varying peripheral blood flow, it is presumed to confer thermoregulatory benefits particularly given the thermal sensitivity of many physiological rate functions in crustaceans.     

Commensalism or mutualism: conditional outcomes in a branchiobdellid–crayfish symbiosis

A growing body of evidence suggests that the costs and benefits involved in cleaning interactions can vary over space and time depending on the environmental conditions. However, whether the same cleaners actually induce variable net outcomes in terms of host fitness remains unclear to date. Branchiobdellid annelids are generally regarded as commensals or rarely parasites of their freshwater crayfish hosts, but a recent study suggests that they may also function as cleaning organisms. Under natural conditions, crayfish can experience fouling of the exposed surfaces of their exoskeletons and their gills (e.g., epibiosis) by various epibionts and particles of organic debris, and branchiobdellids graze on these sources of fouling. Here, we examined the extent to which variation in fouling pressure in the environment alters the outcome of the interaction between branchiobdellids and their crayfish host Cambaroide similis. A series of manipulations were performed in artificial environments designed to simulate either high or low fouling pressure. We used crayfish growth rates and mortality as direct measurements of the net costs and benefits of cleaning. Branchiobdellids had no significant effect on crayfish growth or mortality when cultured under low fouling pressure. However, their presence had a significant positive impact on host growth rates when cultured under high fouling pressure. These results suggest that the relationship between crayfish and branchiobdellids can fluctuate between commensalism and mutualism depending mainly on the environmental fouling pressure. We hypothesize that the outcome of cleaning interactions may largely depend on the factors directly related to the need for cleaning, such as parasite loads or fouling pressure.     

Does juvenile competition explain displacement of a native crayfish by an introduced crayfish?

The coldwater crayfish Orconectes eupunctus is endemic to the Spring and Eleven Point Rivers of Arkansas and Missouri, and appears to have been displaced from a portion of its range by the recently introduced ringed crayfish Orconectes neglectus. We examined competition among juveniles as a potential mechanism for this crayfish species displacement through laboratory and field experiments. Orconectes eupunctus juveniles survived and grew in stream cages in their former range, implicating biotic interactions rather than habitat degradation in the displacement. Laboratory experiments revealed O. neglectus juveniles were dominant in the presence of limited food, whereas size rather than species determined occupancy of limited shelter. In a field competition experiment using stream cages, O. neglectus juveniles did not inhibit growth or reduce survival of O. eupunctus juveniles. Consequently, laboratory evidence of O. neglectus dominance did not correspond with competition under field conditions. Combined with previous studies examining the effects of O. neglectus on O. eupunctus, these results suggest that competition may not be a factor in this crayfish species displacement. Alternate mechanisms for the apparent displacement of O. eupunctus by O. neglectus, such as differential predation or reproductive interference, should be investigated.     

Changes in prostaglandin E2 and F2α during vitellogenesis in the florida crayfish Procambarus paeninsulanus

While the role of eicosanoids in reproduction in vertebrate species has been well established, the role of these fatty acid derivatives in invertebrate species has not been as well characterized. The purpose of this study was to investigate changes in prostaglandins E₂ and F₂ during vitellogenesis in the crayfish Procambarus paeninsulanus. In homogenates of crayfish ovaries taken at various stages of development, the rate of prostaglandin synthesis and the concentrations of prostaglandins E₂ and F₂ increased during the final stages of yolk production just prior to ovulation. A gradual increase in prostaglandin E₂ amounts was observed throughout the progression of vitellogenesis. The data suggests the possible involvement of prostaglandins in regulatory events associated with vitellogenesis and the induction of ovulation in Procambarus paeninsulanus.Abbreviations EIA enzyme immunoassay kit - GIH gonad inhibiting hormone - HPLC high-pressure liquid chromatography - PGE₂ prostaglandin E₂ - PGF₂ prostaglandin F₂ - RIA radioimmuno assay - SNK Student-Newman-Keuls multiple t-test - TLC thin-layer chromatography     

Ontogenetic shift in crayfish δ13C as a measure of land-water ecotonal coupling

Although ontogenetic changes in the carbon isotope ratios of marine fauna have been well studied those of freshwater organisms have not. As a result, we may have a less than adequate assessment of the incorporation of allochthonous detritus into freshwater foodwebs. This study found a ¹³C range of 9 for crayfish (Orconectes virilis) from oligotrophic Canadian Shield lakes. Much (60–83%) of this variability was explained by body size. A simple isotopic mixing model suggests that by their third year of life, crayfish in these lakes rely more substantially upon terrestrial detritus than epilithic algae for energy.     

Branchiobdellid annelids and their crayfish hosts: are they engaged in a cleaning symbiosis?

Branchiobdellid annelids and their freshwater crayfish hosts are generally thought to have a commensal relationship. Branchiobdellids of the genus Cambarincola exploit their hosts through a variety of mechanisms; however, an effect of branchiobdellids on crayfish has not been conclusively demonstrated. We investigated whether branchiobdellids positively affect the host crayfish Cambarus chasmodactylus in the New River, North Carolina. In a laboratory experiment, we placed 0, 3, or 6 branchiobdellids on C. chasmodactylus and observed a significant effect of branchiobdellid presence on both growth and mortality of host crayfish; crayfish with branchiobdellids exhibited faster growth and lower mortality with increasing branchiobdellid density. A tracer experiment demonstrated that branchiobdellids feed on items found in the branchial chamber of C. chasmodactylus. We hypothesize that such feeding activity by branchiobdellids reduces fouling of crayfish gills by epibionts and particulate matter and could lead to the reduced mortality and increased growth rates observed in the laboratory experiment. Specifically, Cambarincola may improve the ventilatory and excretory fitness of C. chasmodactylus by cleaning gill filaments. Field data support this hypothesis by demonstrating that branchiobdellids are found disproportionately at sites proximal to the branchial chamber in the New River. This study provides evidence that the relationship between C. chasmodactylus and Cambarincola may be a cleaning symbiosis, at least in environments where gill fouling is a problem for C. chasmodactylus.     

Do antennule and aesthetasc structure in the crayfish Orconectes virilis correlate with flow habitat?

The local flow environment affects the shape of waterborne chemical signals through a variety of physical mechanisms and at several scales. Since crayfish rely on these chemical signals to extract information about predators, prey, and mates, one might expect the chemical sensors (aesthetascs) on crayfish antennules to be physically tuned to the presentation of chemical cues by the flow environment. This hypothesis was tested by comparing length, diameter, and spacing of antennules and aesthetascs among geographically distinct populations of Orconectes virilis. Crayfish were collected from the Chagrin river, Hebron hatchery, and Burt lake. In addition, antennules were sampled from 43 museum populations representing 12 lake, 10 creek, and 21 river populations from multiple states and river drainages. Mean velocities from the collection sites were either measured directly or calculated from United States Geological Survey (USGS) historical data. Structural parameters were measured using Scion Image software on Scanning electron microscope micrographs, and analyses of variance were performed using StatView. Structural parameters of aesthetascs were found to vary with flow environment. Aesthetascs from lake populations were inserted at a larger angle, extended out farther from the supporting antennule relative to the width of the antennule, and were more widely spaced than aesthetascs from creek, hatchery, and river populations.     

Is inward calcium current in crayfish muscle membrane constituted of one or two components?

Two inward currents were observed in crayfish muscle membrane during depolarization steps by the method described by Adrian et al. (1970). Under voltage clamp conditions, hyperpolarization steps elicited a large current (leak current If), associated with an inward voltage dependent current. This inward current was inhibited by niflumic acid (NA), a drug known to block Cl---HCO-3 exchange (Cousin et Motais 1982; Br?lè et al. 1983b). Dynamic outward currents triggered by depolarizing steps were inhibited to a great extent by TEA, the not inhibited portion disappearing when procaine (2 mmol/l) was added to external solution. In the presence of TEA, procaine and NA, it was thus possible to dissect the regenerative calcium current (ICa) into two components: a "fast component" (ICa1) and a "slow component" (ICa2). The reversal potential of ICa was 65 mV (for [Ca]0 = 2.8 mmol/l), and [Ca]i could be calculated to be 1.6 X 10(-5) mol/l. This value of [Ca]i is the same as calculated from values reported by Hencek and Zachar (1977). ICa1 was triggered at a threshold membrane potential of -45 mV and ICa2 at -30 mV. Moreover, the inactivation kinetics for ICa1 was faster than that for ICa2. Our results are in perfect agreement with those obtained by Zahradník and Zachar (1982) who postulated two populations of calcium channels.     

Fluctuating fecundity parameters and reproductive investment in crayfish: driven by climate or chaos?

1. The striking variation in fecundity of the noble crayfish (Astacus astacus) was assessed from 1981 to 2009 in the lake Steinsfjorden, Norway. 2. Egg size showed no distinct variation with crayfish size or between years and seems to be a fairly fixed property. Ovarian egg numbers increased linearly with body size, from a mean of 70 eggs at maturity to nearly 250 eggs in the largest individuals. There were strong and significant interannual differences in the size‐specific fecundity, and variation in size and among years explained 60% of the observed variance in fecundity. 3. The proportion of mature females in the population ranged from 0.068 to 0.817, with extreme year‐to‐year fluctuations. The proportion of mature females increased significantly over the years, in parallel with increased temperature, but we were unable to link maturity with ambient temperature either in the same year or in the previous year. 4. While several indices point to increased crayfish productivity with higher temperature in this population, there is also a complex and apparently chaotic interplay between life‐cycle attributes, density‐dependent parameters and ambient conditions that lead to high variability in annual recruitment. This probably reflects the slow and delayed life‐cycle responses to a changing environment.     

Do introduced crayfish affect benthic fish in stony littoral habitats of large boreal lakes?

Invasive crayfish are spreading rapidly across Europe, where they are replacing the native crayfish species and impacting negatively on some other biota. Freshwater crayfish and many benthic fishes share similar habitat and food requirements and hence potentially compete for resources. In this study, we investigated impacts of the introduced signal crayfish (Pacifastacus leniusculus) on fish in stony littoral habitats of two large boreal lakes. We compared the littoral fish community composition and the densities of two common benthic fish species between sites with and without crayfish. To evaluate whether signal crayfish share the same food resources as benthic littoral fish or change their feeding habits, we used mixing models and trophic niche estimates based on analyses of stable isotopes of carbon and nitrogen. Both the community composition of littoral fish and the densities of benthic fish species were similar at sites with and without signal crayfish. Even though stable isotope signatures indicated strong dietary overlap between crayfish and benthic fish, the use of food sources and trophic niche widths of fish were not noticeably different between crayfish sites and non-crayfish sites. Our results suggest that, at current densities, the non-native signal crayfish does not have significant impacts on benthic fish in the stony littoral habitats of large boreal lakes.     

Sodium channel inactivation in the crayfish giant axon. Must channels open before inactivating?

Experiments on sodium channel inactivation kinetics were performed on voltage-clamped crayfish giant axons. The primary goal was to investigate whether channels must open before inactivating. Voltage-clamp artifacts were minimized by the use of low-sodium solutions and full series resistance compensation, and the spatial uniformity of the currents was checked with a closely spaced pair of electrodes used to measure local current densities. For membrane potentials between -40 and +40 mV, sodium currents decay to zero with a single exponential time-course. The time constant for decay is a steep function of membrane potential. The time-course of inactivation measured with the double-pulse method is very similar to the decay of current at the same potential. Steady-state inactivation curves measured with different test pulses are identical. The time-course of double pulse inactivation shows a lag that roughly correlates with the opening of sodium channels, but detailed comparisons with the time course of the prepulse current suggest that it is not strictly necessary for channels to open before inactivating. Measurements of the potential dependence of the integral of sodium conductance area also inconsistent with the simplest cases of models in which channels must open before inactivating.     

Invasions and niche width: does niche width of an introduced crayfish differ from a native crayfish?

1. Human activities have promoted the spread of species worldwide. Several crayfish species have been introduced into new areas, posing a threat to native crayfish and other biota. Invader success may depend on the ability to utilise a wide variety of habitats and resources. Successful invaders are generally expected to have broader niches and to be more plastic than non-invasive species.
2. Using stable isotope ratios of carbon and nitrogen we compared the niche widths of native noble crayfish and introduced signal crayfish, a successful invader of Swedish streams. The calculation of niche width took account of between-site differences in basal resource isotope signature ranges. We also assessed whether population density, prey biomass or prey diversity affected niche width.
3. At the species level, signal crayfish had twice the niche width of noble crayfish. However, individual populations of noble crayfish and signal crayfish in Swedish streams had similar niche widths. This suggests that signal crayfish has greater plasticity with respect to habitat utilisation and feeding than noble crayfish. Niche width in both species correlated positively with benthic invertebrate biomass and diversity, indicating that animal food sources are important for crayfish.
4. We find that assessing niche width in relation to invader success can be a useful tool trying to predict the impact of invasions on different scales. The findings in this study suggest that invaders and natives will have a similar impact on the stream scale whereas the invader will have a larger impact on the regional scale due to the ability to utilise a wider range of streams.     

Do rusty crayfish (Faxonius rusticus) invasions affect water clarity in north temperate lakes?

Limnology - Invasive crayfish can cause shifts in lakes from clear, macrophyte-dominated states to eutrophic, phytoplankton-dominated states because of their burrowing and foraging behavior. While...     

Global diversity of crayfish (Astacidae, Cambaridae, and Parastacidae––Decapoda) in freshwater

The freshwater crayfishes are distributed across all but the Indian and Antarctic continents with centers of diversity in the southeastern Appalachian Mountains in the Northern Hemisphere and in south–east Australia in the Southern Hemisphere. There are currently over 640 described species of freshwater crayfishes with an average of 5–10 species still being described each year. Freshwater crayfishes can serve as keystone species in aquatic habitats, but a few species are also significantly invasive and can cause impressive damage to the fragile freshwater habitat. Crayfishes inhabit caves, burrows, streams, lakes and strong burrowers can even be found in terrestrial habitats where they have burrowed to the water table or where rainfall is sufficiently abundant to provide the needed moisture. The freshwater crayfishes, like the habitats in which they are encountered, are generally endangered to some degree and conservation efforts would do well to focus on them as key elements of the freshwater ecosystem. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Are fish populations in temperate streams affected by crayfish? – A field survey and prospects

Synopsis Fish populations may be affected by predation and competition from various types of organisms, among which crayfish have been suggested as important actors. We here present results from stream surveys, suggesting that neither native noble, Astacus astacus, nor introduced signal crayfish, Pacifastacus leniusculus, necessarily affect fish population densities in temperate stream communities. Comparisons of fish densities within stream sites between years with absence and presence of crayfish showed no effect of either crayfish species. A further analysis of changes in fish densities between periods without and with crayfish in low, intermediate and high densities revealed that crayfish density did neither have an effect on fish densities. Our study is one of exceptionally few that consider the above aspects in long-term perspectives in natural systems, and we discuss that previously reported divergent results of crayfish effects on fish may be highly dependent on specific species and methods used, and that the effects of crayfish on fish populations deserve further attention to enable reliable predictions of community processes in streams.