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.     

Over-invasion in a freshwater ecosystem: newly introduced virile crayfish (Orconectes virilis) outcompete established invasive signal crayfish (Pacifastacus leniusculus)

Biological invasions are a key threat to freshwater biodiversity, and identifying determinants of invasion success is a global conservation priority. The establishment of introduced species is predicted to be hindered by pre-existing, functionally similar invasive species. Over a five-year period we, however, find that in the River Lee (UK), recently introduced non-native virile crayfish (Orconectes virilis) increased in range and abundance, despite the presence of established alien signal crayfish (Pacifastacus leniusculus). In regions of sympatry, virile crayfish had a detrimental effect on signal crayfish abundance but not vice versa. Competition experiments revealed that virile crayfish were more aggressive than signal crayfish and outcompeted them for shelter. Together, these results provide early evidence for the potential over-invasion of signal crayfish by competitively dominant virile crayfish. Based on our results and the limited distribution of virile crayfish in Europe, we recommend that efforts to contain them within the Lee catchment be implemented immediately.     

Niche differentiation among invasive crayfish and their impacts on ecosystem structure and functioning

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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.     

Functional distance and establishment of non-native species with complex life cycles

More than 80% of animals have complex life cycles and undergo distinct changes in ecology and morphology during development. The strength and type of factors regulating each life-stage may differ as an organism may occupy different niches during ontogeny. We examined the functional distance at larval and adult life-stages of two non-native anurans (Green Tree Frog [Hyla cinerea] and Bullfrog [Lithobates catesbeianus]) that have established in a Chihuahuan Desert anuran assemblage in Big Bend National Park. Both life stages of both non-native species occupied niche space outside of the native assemblage. At the larval stage, the ability of the tadpoles to utilize permanent aquatic habitats and coexist with predatory fishes differentiated the non-native species from the majority of the native species that are restricted to temporary pools. At the post-metamorphic life stage, each species appears to have established by exploiting unoccupied habitat and trophic niches in the recipient community. The arboreal habits of H. cinerea may enable it to utilize resources in microhabitats that are otherwise not used by native species because arboreal frogs are absent from this native assemblage. The large body size of post-metamorphic L. catesbeianus may enable it to utilize larger food resources that are otherwise unavailable to the smaller-bodied natives. Separate comparison of larval and adult functional traits between non-natives and the native community may help predict their potential establishment or invasion success as well as aid in the development of stage-specific control or eradication efforts.     

Associations between stream habitat characteristics and native and alien crayfish occurrence

Human mediated introductions of non-indigenous crayfish species (NICS) are responsible for their rapid colonisation of European freshwaters. The introduction of North American crayfish is furthermore linked to the spread of crayfish plague and the decline of indigenous crayfish species (ICS). As the management of ICS and NICS have become necessary, a detailed knowledge on their distribution and ecological requirements is needed. We studied the current range of native noble crayfish Astacus astacus and stone crayfish Austropotamobius torrentium, as well as alien signal crayfish Pacifastacus leniusculus in Carinthia (Austria) and evaluated environmental and physical habitat features in streams with and without crayfish. Meanwhile, the loss of many ICS populations was recorded and alien P. leniusculus was found to be widespread in this region. Most of the habitat features of streams having crayfish differed significantly from sites lacking crayfish for at least one investigated native or alien species. Furthermore, multivariate and regression analyses showed specific differences in the habitat use of the investigated crayfish. Our results showed that the presence of alien P. leniusculus was associated with larger and smoother sloped lowland rivers, while the occurrence of the two native species was confined to smaller streams either at higher altitudes and with distinct physical habitat conditions (A. torrentium) or with moderate water temperatures (A. astacus). This study helps to identify potential refuge areas for the endangered native species and to predict the further spread of the most common non-native crayfish species in European streams.     

Invasive pythons,not anthropogenic stressors,explain the distribution of a keystone species

Untangling the causes of native species loss in human-modified systems is difficult and often controversial. Evaluating the impact of non-native species in these systems is particularly challenging, as additional human perturbations often precede or accompany introductions. One example is the ongoing debate over whether mammal declines within Everglades National Park (ENP) were caused by either the establishment of non-native Burmese pythons (Python molurus bivittatus) or the effects of other anthropogenic stressors. We examined the influence of both pythons and a host of alternative stressors—altered hydrology and habitat characteristics, mercury contamination and development—on the distribution of the marsh rabbit (Sylvilagus palustris), a once common mammal in ENP. Distance from the epicenter of the python invasion best explained marsh rabbit occurrence in suitable habitat patches, whereas none of the alternative stressors considered could explain marsh rabbit distribution. Estimates of the probability of marsh rabbit occurrence ranged from 0 at the python invasion epicenter to nearly 1.0 150 km from the invasion epicenter. These results support the hypothesis that invasive pythons shape the distribution of marsh rabbits in southern Florida. The loss of marsh rabbits and similar species will likely alter trophic interactions and ecosystem function within the Everglades, an internationally important hotspot of biodiversity. Further, our results suggest that non-native species can have profound impacts on mainland biodiversity.     

Geographic independence and phylogenetic diversity of red shiner introductions

Identifying areas at risk of invasion can be difficult when the distribution of a non-native species encompasses geographically disjunct regions. Understanding genealogical relationships among native and non-native populations can clarify the origins of fragmented distributions, which in turn can clarify how fast and far a non-native species may spread. We evaluated genetic variation across the native and invasive ranges of red shiner (Cyprinella lutrensis), a minnow known to displace and hybridize with native species, to reconstruct invasion pathways across the United States (USA). Examination of mitochondrial cytochrome-b variation found that native range populations of red shiner fall into four highly divergent lineages that likely warrant species recognition. Introduced red shiner populations in the eastern and western USA are derived from only two of these lineages. Western USA populations originate from the mid-western and western genetic lineages, whereas eastern introductions derive only from the mid-western lineage. Western USA invasive populations exhibit fewer, but more diverse haplotypes compared to eastern USA invasive populations. We also recovered an undescribed, divergent lineage of Cyprinella that has been cryptically introduced into the western USA, which raises the possibility that hybridization has proceeded following secondary contact between previously allopatric lineages. Approximate Bayesian Computation modeling suggests that the disjunct distribution of red shiner across North America is an agglomeration of independent regional invasions with distinct origins, rather than stepwise advance of an invasion front or secondary introductions across regions. Thus localized control may be effective in managing non-native red shiner, including further spread to areas of conservation concern.     

Trophic consequences of non-native pumpkinseed <Emphasis Type="Italic">Lepomis gibbosus</Emphasis> for native pond fishes

Introduced non-native fishes can cause considerable adverse impacts on freshwater ecosystems. The pumpkinseed Lepomis gibbosus, a North American centrarchid, is one of the most widely distributed non-native fishes in Europe, having established self-sustaining populations in at least 28 countries, including the U.K. where it is predicted to become invasive under warmer climate conditions. To predict the consequences of increased invasiveness, a field experiment was completed over a summer period using a Control comprising of an assemblage of native fishes of known starting abundance and a Treatment using the same assemblage but with elevated L. gibbosus densities. The trophic consequences of L. gibbosus invasion were assessed with stable isotope analysis and associated metrics including the isotopic niche, measured as standard ellipse area. The isotopic niches of native gudgeon Gobio gobio and roach Rutilus rutilus overlapped substantially with that of non-native L. gibbosus, and were also substantially reduced in size compared to ponds where L. gibbosus were absent. This suggests these native fishes shifted to a more specialized diet in L. gibbosus presence. Both of these native fishes also demonstrated a concomitant and significant reduction in their trophic position in L. gibbosus presence, with a significant decrease also evident in the somatic growth rate and body condition of G. gobio. Thus, there were marked changes detected in the isotopic ecology and growth rates of the native fish in the presence of non-native L. gibbosus. The implications of these results for present and future invaded pond communities are discussed.     

Phenotypic variability of rusty crayfish (Faxonius rusticus) at the leading edge of its riverine invasion

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Invasive and Non-Invasive Congeners Show Similar Trait Shifts between Their Same Native and Non-Native Ranges

Differences in morphological or ecological traits expressed by exotic species between their native and non-native ranges are often interpreted as evidence for adaptation to new conditions in the non-native ranges. In turn this adaptation is often hypothesized to contribute to the successful invasion of these species. There is good evidence for rapid evolution by many exotic invasives, but the extent to which these evolutionary changes actually drive invasiveness is unclear. One approach to resolving the relationship between adaptive responses and successful invasion is to compare traits between populations from the native and non-native ranges for both exotic invaders and congeners that are exotic but not invasive. We compared a suite of morphological traits that are commonly tested in the literature in the context of invasion for three very closely related species of Centaurea, all of which are sympatric in the same native and non-native ranges in Europe and North America. Of these, C. solstitialis is highly invasive whereas C. calcitrapa and C. sulphurea are not. For all three species, plants from non-native populations showed similar shifts in key traits that have been identified in other studies as important putative adaptive responses to post-introduction invasion. For example, for all three species plants from populations in non-native ranges were (i) larger and (ii) produced seeds that germinated at higher rates. In fact, the non-invasive C. calcitrapa showed the strongest trait shift between ranges. Centaurea solstitialis was the only species for which plants from the non-native range increased allocation to defensive spines, and allocated proportionally less resources to reproduction, patterns contrary to what would be predicted by theory and other empirical studies to enhance invasion. Our results suggest caution when interpreting the commonly observed increase in size and reproductive capacity as factors that cause exotics to become invaders.     

Ecological niche differentiation between native and non-native shrimps in the northern Baltic Sea

Invasions of non-native species are modifying global biodiversity but the ecological mechanisms underlying invasion processes are still not well understood. A degree of niche separation of non-native and sympatric native species can possibly explain the success of novel species in their new environment. In this study, we quantified experimentally and in situ the environmental niche space of caridean shrimps (native Crangon crangon and Palaemon adspersus, non-native Palaemon elegans) inhabiting the northern Baltic Sea. Field studies showed that the non-native P. elegans had wider geographical range compared to native species although the level of habitat specialization was similar in both Palaemon species. There were clear differences in shrimp habitat occupancy with P. elegans inhabiting lower salinity areas and more eutrophicated habitats compared to the native species. Consequently, the non-native shrimp has occupied large areas of the northern Baltic Sea that were previously devoid of the native shrimps. Experiments demonstrated that the non-native shrimp had higher affinity to vegetated substrates compared to native species. The study suggests that the abilities of the non-native shrimp to thrive in more stressful habitats (lower salinity, higher eutrophication), that are sub-optimal for native shrimps, plausibly explain the invasion success of P. elegans.     

Comparison of trophic function between the globally invasive crayfishes <Emphasis Type="Italic">Pacifastacus leniusculus</Emphasis> and <Emphasis Type="Italic">Procambarus clarkii</Emphasis>

Impacts of invasive species may manifest most strongly if these organisms are highly distinct functionally from the native species they often replace. Yet, should we expect functional differences between native and invasive species of generalist organisms like freshwater crayfish? Some existing evidence has pointed to native and invasive crayfish species as ecologically equivalent. We contribute to this literature by comparing the trophic niches of the globally invasive crayfishes Pacifastacus leniusculus and Procambarus clarkii, by applying carbon and nitrogen stable isotope analyses to replicated allopatric (alone) and sympatric (together) lake populations in western Washington State, USA, where P. clarkii has been recently introduced and P. leniusculus is presumed native. Our study corrected for potential inherent differences in lake food webs as a consequence of lake abiotic or biotic characteristics using random effects in linear mixed effects models. We found that although overall trophic niche size or area of these species was not significantly different, P. leniusculus was significantly higher in trophic position than P. clarkii when also accounting for the effects of body size, sex, and lakes as random effects. This pattern of increased trophic position of P. leniusculus over P. clarkii was conserved over time in one sympatric lake for which we had data over multiple years. Cumulatively, our findings point to trophic differences between the globally cosmopolitan crayfishes P. leniusculus and P. clarkii, particularly when accounting for the ways that ecosystem context can affect food web structure of communities and the trophic resources available to these consumers.     

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.     

Non-native aquatic animals introductions have driven disease emergence in Europe

In this paper it is argued, using examples of disease emergence in aquatic animals in Europe, that the introduction of non-native species drives disease emergence by both extending the geographic range of parasites and pathogens and facilitating host-switching. Enteric red mouth disease and infectious haematopoietic necrosis of salmonids have extended their geographic range from North America to Europe with the import of live fish (Pimephales promelas) and rainbow trout eggs, respectively. Host-switching results in disease emergence when previously unidentified commensal organisms or known pathogen switch to new naïve hosts. The most serious endemic diseases of wild aquatic animals in Europe in recent years can be traced to the introduction of non-native species. Across Europe dramatic populations declines have occurred in native crayfish (e.g. Astacus astacus), oysters (Ostrea edulis) and eels (Anguilla anguilla), all which can be attributed, in varying degrees, to diseases (crayfish plague, Bonamia ostreae and Anguillicoloides crassus, respectively) introduced with non-native species. The severe adverse effects at a population level can be attributed to the lack of immunity in the new hosts. The impact of parasites more recently introduced to Europe, Sphaerothecum destruens (the rosette agent), and Batrachochytrium dendrobatidis, have yet to be fully determined. Both are generalists, with wide host ranges, and may present serious threats to native species. Aquaculture is the key driver for the introduction of non-native species. Most farming systems allow pathogen exchange between farmed and wild populations which underpins host-switching. Subsequently movements of animals between farms may result in the spread of newly emerged diseases. The introduction of non-native aquatic animals drives disease emergence, thus the ex-ante assessment of these hazards is severely limited. Generic risk mitigation measures (e.g. movement of disinfected eggs in place of live animals) and improved methods for rapid detection of new diseases are vital.     

入侵植物的繁殖策略以及对本土植物繁殖的影响

理解入侵生物的繁殖策略是阐明生物入侵机制的一个重要方面。入侵植物常表现出一些共同的繁殖特征, 如以两性花为主的性系统、自动自交为主的繁育系统或不依赖传粉媒介的无融合生殖和无性繁殖以及高生殖投资的资源配置策略等。成功入侵的外来植物通过影响本土的传粉者, 在种群和群落水平上影响本土植物的有性繁殖, 甚至促使某些本土植物在繁殖对策和表型性状上发生快速转变。目前, 入侵植物繁殖策略及其生态效应的研究多侧重于入侵种的快速演化, 而有关外来植物与本土植物间的相互影响及其可能存在的协同适应研究还较为缺乏。探讨本土植物在外来种入侵压力下的繁殖对策和响应机制, 将丰富人们对物种间竞争、共存及群落构建等机制的深入了解。从繁殖和适应的角度探求入侵植物与本土植物之间的复杂关系, 将有助于解析生物入侵的机制及人类干扰下的物种演化规律, 也为预测和防控入侵植物提供科学依据。     

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.     

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.     

Transformation of detritus by a European native and two invasive alien freshwater decapods

Invasive alien species have the potential to alter biodiversity and ecosystem processes. In freshwaters, detritus decomposition is a major ecosystem service but it remains uncertain whether invasive alien decapods process detritus differently to natives. This study examined leaf litter processing, and cascading effects on biofilms, by the European native white clawed crayfish (Austropotamobius pallipes) compared to two invasive alien decapod species: the American signal crayfish (Pacifastacus leniusculus) and the Chinese mitten crab (Eriocheir sinensis). Invasive alien decapods were responsible for higher leaf litter decomposition than the native. In comparison with native crayfish, invasive alien crab and crayfish showed higher rates of litter consumption, increased production of smaller leaf fragments, fine particulate organic matter (FPOM) and dissolved organic carbon. Nutrients (ammonia and soluble reactive phosphorous) derived from excretion (measured separately in the absence of biofilms) varied among decapod species, being lower for P. leniusculus. However, nutrient concentrations did not vary among species in the detritivory experiments with biofilm, implying nutrients were utilised for biofilm production and respiration as no differences in biomass were evident among decapod treatments. These results show invasive alien decapods have the potential to increase the magnitude of detrital processing to FPOM in rivers, but indirect impacts on primary producers due to nutrient release are uncertain based on this experimental context.     

Let’s eat out,there’s crayfish for dinner: American bullfrog niche shifts inside and outside native ranges and the effect of introduced crayfish

In invasion ecology, niche width has been recognized as a crucial factor for the outcome of an invasion. A common characteristic of successful invaders seems to be a broad niche width, and their impact on native communities may increase with increasing niche size. Overall, successful invader predators are predicted to shift their niche width by broadening it from native to invaded conditions. The scarcity of ecological studies examining invasive species in their native ranges prevents researchers from knowing if the prevalence of generalist invaders represents conservatism of broad native-range niches or instead niche shifts as a result of different processes acting in the invaded areas. Here we reviewed literature on trophic niche of the predatory invader American bullfrog (Lithobates catesbeianus) in both native and invaded ranges. We used statistical and graphic tools to analyse possible shifts in dietary niche width and the effect of introduced crayfish on the feeding strategy of L. catesbeianus. Globally, our results indicate that food sources used by the species differed in native and invaded sites, with a narrower trophic niche width in invaded areas. However, this pattern was disrupted by the occurrence of introduced crayfish that represents the major driver of the observed niche-width variation. Our data shed light on possible complications in interpreting and predicting patterns of biological invasions due to the interaction among species from different trophic levels that apparently disrupt general patterns that are likely bound to be idiosyncratic and complex.