High niche overlap between native and non-native apex fish in a human-impacted floodplain

To test ecological niche theory, this study investigated the spatial patterns and the environmental niches of native and non-native fishes within the invaded Great Fish River system, South Africa. For the native fishes, there were contrasting environmental niche breadths that varied from being small to being large and overlapped for most species, except minnows that were restricted to headwater tributaries. In addition, there was high niche overlap in habitat association among fishes with similar distribution. It was therefore inferred that habitat filtering-driven spatial organisation was important in explaining native species distribution patterns. In comparison, most non-native fishes were found to have broad environmental niches and these fishes showed high tolerance to environmental conditions, which generally supported the niche opportunity hypothesis. The proliferation of multiple non-native fishes in the mainstem section suggest that they form a functional assemblage that is probably facilitated by the anthropogenic modification of flow regimes through inter-basin water transfer. Based on the distribution patterns observed in the study, it was inferred that there was a likelihood of negative interactions between native and non-native fishes. Such effects are likely to be exacerbated by altered flow regime that was likely to have negative implications for native ichthyofauna.

Pilger TJ, Gido KB, Propst DL. Diet and trophic niche overlap of native and nonnative fishes in the Gila River, USA: implications for native fish conservation. Ecology of Freshwater Fish 2010: 19: 300–321. © 2010 John Wiley & Sons A/SAbstract – The upper Gila River basin is one of the few unimpounded drainage basins west of the Continental Divide, and as such is a stronghold for endemic fishes in the region. Nevertheless, multiple nonindigenous fishes potentially threaten the persistence of native fishes, and little is known of the trophic ecology of either native or nonnative fishes in this system. Gut contents and stable isotopes (13C and 15N) were used to identify trophic relationships, trophic niche overlap and evaluate potential interactions among native and nonnative fishes. Both native and nonnative fishes fed across multiple trophic levels. In general, adult native suckers had lower 15N signatures and consumed more algae and detritus than smaller native fish, including juvenile suckers. Adult nonnative smallmouth bass (Micropterus dolomieu), yellow bullhead (Ameiurus natalis) and two species of trout preyed on small‐bodied fishes and predaceous aquatic invertebrates leading to significantly higher trophic positions than small and large‐bodied native fishes. Thus, the presence of these nonnative fishes extended community food‐chain lengths by foraging at higher trophic levels. Although predation on juvenile native fishes might threaten persistence of native fishes, the high degree of omnivory suggests that impacts of nonnative predators may be lessened and dependent on environmental variability.

Populations of many native freshwater mussels such as the California floater, Anodonta californiensis, are declining in the Columbia River Basin in the western United States. There are many possible reasons for this decline, including increased presence of nonnative fishes, especially piscivores that displace and reduce the abundance of native fishes. These nonnative fishes can negatively influence the early life history of A. californiensis, because this mussel uses native fishes to complete its life cycle. While Anodonta spp. can use nonnative fishes as hosts with limited success, the extent of this for A. californiensis is not well understood. We determined if the glochidia (larvae) of A. californiensis can successfully metamorphose using certain nonnative fishes, and we quantified differences in host effectiveness (number of juveniles produced) between native and nonnative fishes based on available fin area for attachment. Overall, native fishes hosted an average of 107.4 ± 39.9 (mean ± SE) juvenile mussels per fish while nonnative fishes hosted an average of 5.5 ± 4.9 juveniles per fish. This conclusion was unchanged when standardized for fish attachment area. Encystment of glochidia on native fishes yielded an average of 1.0 ± 0.1 juveniles/mm2 of attachable surface (total area of fins) area while nonnative fishes yielded an average of 0.16 ± 0.1 juveniles/mm2. The nonnative channel catfish, Ictalurus punctatus, did not yield any juvenile mussels and was identified as a nonhost species for A. californiensis. All other nonnative fishes tested were determined to be poor or marginal hosts. All native fishes were determined to be primary or secondary hosts for A. californiensis. The native fishes that yielded the highest number of juvenile mussels were sculpins, Cottus spp., and the redside shiner, Richardsonius balteatus, with an average of 196 and 151 juveniles per individual of each species, respectively. Our findings show that nonnative fishes are poor hosts for A. californiensis. However, nonnative fishes may contribute to the decline of native mussels in the Columbia River Basin by directly preying on and reducing the abundance of native host fishes and mussels. Future conservation plans for A. californiensis should consider the potential negative influence of nonnative fishes.

Non-native fishes homogenize native fish communities and reduce ecosystem multifunctionality in tropical lakes over 16 years

Aquatic ecosystems can harbour more than one non-native fish species and this can represent a threat due to trophic interactions with native fishes. However, research on interactions amongst multiple co-occurring native and non-native fish remains scarce. In this study, 551 organisms from 44 native fish, 11 non-native fish, 35 macroinvertebrates (of which one was non-native), together with 162 samples of basal resources were collected from six rivers of the Lower Pearl River Basin of China. Nitrogen and carbon stable isotope analysis was used to calculate community-wide trophic metrics and the degree of trophic overlap between native and non-native fish at both the community and functional feeding group level, together with diet composition. At the community level, there was a high degree of trophic niche overlap between native and non-native fish as a result of similarities in trophic characteristics. At the functional feeding group level, both native and non-native functional feeding groups demonstrated the capacity to occupy the niche space of each other. A significant trophic niche overlap, exceeding 50%, was found between non-native detritivorous and omnivorous fish, suggesting competition. The difference in diet composition between some native and non-native fish depended on the category of diet source across the rivers, suggesting dietary segregation. Albeit limited, the present findings suggest that trophic interaction between native and non-native fish is likely to reach a dynamic equilibrium status in the community owing to trophic segregation of fish species and the antagonistic effects amongst non-native fish.

The strength of the interspecific interactions between non-native and native species influences the subsequent invasion impacts, with stronger interactions and thus more severe impacts predicted when the species are phylogenetically close and co-exist at lower latitudes. Although work demonstrates non-native fish are more likely to invade areas which have phylogenetically closely related species present, the impacts of these invaders on the native species remain poorly understood. Accordingly, we conducted a comprehensive analysis of global freshwater fish occurrence data to test the phylogenetic niche conservatism hypothesis in co-occurring native and non-native freshwater fishes, assess whether non-native fishes drive extinctions of phylogenetically closely related native species, and evaluate how this varies with latitude. Phylogenetic niche conservatism was evident in co-occurring non-native and native fishes, with their morphometric trait dissimilarity significantly correlated with their phylogenetic distance. Extinct freshwater fishes were found to be more closely related to co-occurred non-native species than were species of other conservation statuses. However, the relationship between the extinction probability of native freshwater fishes and their phylogenetic relatedness to non-native species was context-dependent, varying with latitude and across biogeographical realms. At higher latitudes, native fishes closely related to non-native species had a higher probability of extinction, whereas at lower latitudes, closely related native species were less likely to become extinct. Thus, the extinction risks posed by non-native fish vary spatially and depend on their phylogenetic relatedness to native species, both of which are recommended for consideration in invasion risk management programmes.

The maintenance or restoration of natural flow regimes has been proposed as one means of conserving native fishes. Native fish conservation is enhanced either through the restoration of natural fluvial geomorphic processes (and thus the maintenance of essential habitats) or by the suppression of nonnative fishes. The San Juan River of Colorado, New Mexico, and Utah was dammed in 1962 and its natural flow regime was lost. Beginning in 1993, the river was regulated to mimic a natural flow regime by increasing reservoir releases to mimic timing, but only partially to mimic amplitude, volume, and duration of spring snowmelt discharge. We evaluated the responses of native and nonnative fishes to this natural flow regime mimicry by comparing their autumn densities (number/m2) in San Juan River secondary channels to those during spring runoff and summer base flow over a 9‐year period. Densities of native speckled dace Rhinichthys osculus, bluehead sucker Catostomus discobolus, and flannelmouth sucker C. latipinnis increased with elevated spring discharge. Total native fish density was 10 times greater in 1993 (the year of highest spring discharge) than in 2000 (the year of lowest spring discharge). Collectively, nonnative fish density was negatively related to spring discharge, but western mosquitofish Gambusia affinis was the only commonly collected nonnative that had a significant relationship. Mean daily summer discharge did not affect the density of native or nonnative fishes. Nonnative fishes, however, responded positively to sustained low summer flows (days discharge was less than 14 m3/s). Densities of red shiner Cyprinella lutrensis, common carp Cyprinus carpio, and western mosquitofish were four or more times greater in 2000 (a year of sustained low summer discharge) than in years with comparatively high summer discharge. Speckled dace was the only native species negatively affected by extended low summer discharge. Our results suggest that manipulating spring discharge to mimic a natural flow regime enhances native fish recruitment but might have limited effect in suppressing nonnative fishes, particularly fecund, rapidly growing, small‐bodied species.

Pool refuges are critical for maintaining stream fish diversity in increasingly intermittent streams. Yet, the patterns and drivers of beta diversity of native and non-native fish in pool refuges remain poorly known. Focusing on Mediterranean streams, we decomposed beta diversity of native and non-native fish into richness difference (RichDiff) and species replacement (Repl), and local (LCBD, LCBDRichDiff and LCBDRepl) and species (SCBD) contributions. We assessed the influence of environmental and spatial factors associated with drought and damming fragmentations on beta diversity components and LCBDs, and of local species richness and occupancy on LCBDs and SCBD, respectively. Overall, non-native species showed a more limited occupancy of pool refuges than native fish. RichDiff dominated beta diversity, though it was influenced by drought and damming fragmentations for native fish and local environment for non-native fish. Repl for native fish was slightly influenced by local environment, but for non-native fish was largely driven by drought and damming, albeit with a contribution of local environment as well. LCBD and LCBDRichDiff increased in pools in low order streams for native fish and at low elevations for non-native fish, and with high or low species richness. SCBD was higher for native species with intermediated pool occupancy, but for non-native species with low occupancy. Our results suggest that stream fragmentation may drive native species loss and non-native species replacement in pool refuges, and that environmental filtering may shape non-native species loss. Pools in lower order streams harbouring unique species-rich or species-poor assemblages should be prioritize for conservation and restoration, respectively, and pools at low elevation with unique non-native assemblages should deserve control efforts. We encourage the partitioning of beta diversity and individual analysis of native and non-native fish in intermittent streams, which may be key in stressing the importance of pool refuges in safeguarding native fish diversity.

Nonnative fishes have been linked to the decline of native fishes and may affect aquatic food webs through direct and indirect pathways. These concerns have led to efforts to remove nonnative Brown and Rainbow Trout, which are abundant in tributaries of the Colorado River, to enhance native fish communities. We sampled fish, benthic, and drifting macroinvertebrates in November 2010, January 2011, June 2011, and September 2011 to assess resource availability and to evaluate the effects of nonnative Brown and Rainbow Trout in a tributary of the Colorado River in the Grand Canyon. We evaluated trout diets from stomach samples collected during macroinvertebrate sampling periods, and we estimated annual consumption with bioenergetics models. We used 13C and 15N stable isotopes to examine potential diet overlap between native and nonnative fishes. Contributions to benthic biomass varied among megalopterans (16–35%), trichopterans (19–28%), and ephemeropterans (9–32%), whereas ephemeropterans dominated biomass (44–64%) in drift samples. Ephemeropterans were dominant in diets of small (<150 mm total length [TL]) trout, whereas Corydalus and native fish dominated diets of large (>150 mm TL) Brown Trout, and Corydalus and algae dominated diets of large Rainbow Trout. Annual resource consumption was 6× higher for large trout than small trout. Stable isotopes suggested diet overlap between native and nonnative fishes. Large nonnative trout occupied the highest trophic positions. Our results suggest that suppression of nonnative trout may have a positive effect on native fishes via reduced predation and resource competition.

Effects of Flooding on Recruitment of Roundtail Chub, Gila robusta, in a Southwestern River

Globally, there is growing concern on the occurrence of multiple non-native species within invaded habitats. Proliferation of multiple non-native species together with anthropogenic-driven habitat modifications raise questions on the mechanisms facilitating the co-occurrence of these species and their potential impact within the recipient systems. Using the Great Fish River system (South Africa) which is anthropogenically-modified by inter-basin water transfer (IBWT), as a case study, this research employed trait-based approaches to explore patterns associated with the co-occurrence of multiple non-native fish species. This was achieved by investigating the role of functional diversity of non-native and native fishes in relation to their composition, distribution and environmental relationships. Nineteen functional traits that defined two broad ecological attributes (habitat use and feeding) were determined for 13 fish species that comprised eight native and five non-native fishes. We used these data to, firstly, evaluate functional diversity patterns and to compare functional traits of native and non-native fishes in the Great Fish River system. Secondly, we employed multivariate ordination analyses (factor analysis, RLQ and fourth-corner analyses) to investigate interspecific trait variations and potential species-trait-environmental relationships. From a functional diversity perspective, there were no significant differences in most functional diversity indices between native and non-native species. Despite interspecific variation in body morphology-related traits, we also found no clear separation between native and non-native species based on the ordination analysis of the functional traits. Furthermore, while RLQ ordination showed broad spatial patterns, the fourth-corner analyses revealed no significant relationships among species distribution, functional traits and environmental variables. The weak species-trait-environment relationship observed in this study suggests that environmental filtering was likely a poor determinant of functional trait structure within the Great Fish River. Modification of the natural flow regime may have weakened the relationship between species traits and the environment as has been shown in other systems.

Invasive fish keeps native feeding strategy despite high niche overlap with a congener species

Simple SummaryInvasive species are widely recognized to negatively affect native species through both direct and indirect interactions. When diet overlap between the native and invasive species increases, their competitive interaction is expected to increase too. This in turn may lead to displacement of one of the species. However, the specific mechanisms of the diet displacement are still unclear. In this study, we analysed the diet and diet overlap between the critically endangered European mink and the invasive American mink during the invasion process of the latter species by means of stable isotope analyses. We found a significant diet overlap between the native and invasive mink when they co-occur, an important individual variation of diet, and no significant change of diet of the native species in response to the arrival of the invasive mink. These results suggest significant competitive pressure imposed on the native European mink by the invasive American mink. As such, urgent implementation of control measures of invasive species is needed to ensure the viability and conservation of endangered European mink populations.The pressure elicited by invasive species on native species significantly increases with the increase of the overlap of their ecological niches. Still, the specific mechanisms of the trophic displacement of native species during the invasion process are unclear. The effects of the invasive American mink (Neovison vison) on the critically endangered European mink (Mustela lutreola) was assessed by analyses of diet and niche overlap during the invasion process. To do this, the isotopic composition (δ13C and δ15N) of both species of mink and their four main types of prey was analysed. Significant trophic overlap between the native European mink and invasive American mink was found when they coexisted in sympatry. Furthermore, both mink species were characterised by significant individual variation in diet and no obvious change in diet of the native species in response to the arrival of the introduced species was observed. High niche overlap registered between both species in sympatry with no displacement in diet of the native mink in response to the arrival of the invasive mink is expected to have important consequences for the viability and conservation of the native mink populations, as it suggests high competitive pressure.

If interspecific competition is a strong structuring force of communities, ecologically similar species should tend to have spatial ranges at local scale that do not overlap. Experimental testing of this hypothesis becomes impracticable with large communities. One possibility to tackle this issue is a correlational approach, by comparing the matrix of niche overlap with that of spatial overlap. The use of the standard Mantel test is however impaired by the non‐linearity in the relationship of the two descriptors: in a competitively structured assemblage, species with high niche overlap are expected to be segregated spatially, but species with small niche overlap may or may not exhibit high spatial overlap. To overcome this problem, we devised an original randomization test, which was run for three data‐sets comprising frogs, lizards, and birds along altitudinal gradients. The test yielded intriguing results: reptiles and birds revealed an adjustment that would reduce the potential for interspecific competition, while amphibians showed the opposite trend, that is, ecologically similar species co‐occurred more than expected by chance. Frogs may be more constrained by resource requirements, possibly breeding sites, than by competition. Our test will help to assess the generality of this pattern with other data‐sets.

Food-web investigations inform management strategies by exposing potential interactions between native and nonnative species and anticipating likely outcomes associated with species removal efforts. We leveraged a natural gradient of compositional turnover from native-only to nonnative-only fish assemblages, combined with an intensive removal effort, to investigate underlying food-web changes in response to invasive species expansion in a Lower Colorado River tributary. Nonnative fishes caused coordinated isotopic niche displacement in native fishes by inducing resource shifts toward lower trophic positions and enriched carbon sources. By contrast, nonnative fishes did not experience reciprocal shifts when native fishes were present. Asymmetrical outcomes between native and nonnative fishes indicated species displacement may result from competitive or consumptive interactions. Native species’ isotopic niches returned to higher trophic levels after nonnative green sunfish (Lepomis cyanellus) removal, indicating removal efforts can support trophic recovery of native fishes like desert suckers (Catostomus clarkii) and roundtail chub (Gila robusta). Using stable isotope analysis in preremoval assessments provides opportunities to identify asymmetric interactions, whereas postremoval assessments could identify unintended consequences, like mesopredator release, as part of adaptive decision making to recover native fishes.