Factors affecting the establishment success of the invasive piscivorous chub in small irrigation ditches in northern Kyushu,Japan

Habitat matching is an important factor for the establishment of alien fishes in recipient environments. The piscivorous chub, Opsariichthys uncirostris uncirostris, originally exclusively inhabited large water areas; however, the species has been introduced and established in small irrigation ditches on Kyushu Island. The evaluation of the habitat use of the piscivorous chub suggested that this alien fish uses the lentic-type irrigation ditches as a nursery ground and the lotic-type ditches as a spawning ground. This finding implies that the establishment of the piscivorous chub may require connectivity between lentic and lotic habitats.     

Predicting the spatial distribution of the invasive piscivorous chub (<Emphasis Type="Italic">Opsariichthys uncirostris uncirostris</Emphasis>) in the irrigation ditches of Kyushu,Japan: a tool for the risk management of biological invasions

The piscivorous chub (Opsariichthys uncirostris uncirostris) has widely invaded Kyushu Island in Japan, and its presence in irrigation ditches known as creeks around Ariake Bay has caused particular concern because various native freshwater fishes are also known to exist in the region. In order to examine the habitat characteristics that are related to its occurrence, we developed a species distribution model for piscivorous chub that inhabits creeks in the Kase river catchment by using geographic and habitat variables that were both biotic and abiotic. We then evaluated the model by using a different data set from the adjacent Chikugo river catchment. The resulting multiple logistic regression model, whose performance was supported by a high value of 0.881 for the area under the receiver operating characteristics curve (AUC), indicated that the occurrence of piscivorous chub was strongly affected by the watercourse distance from the source populations in the Kase river. The model’s performance was still high (AUC = 0.792) when tested with the data set from the Chikugo river catchment. We also produced a GIS map that projects the predicted distribution of piscivorous chub across all creeks within the Kase river catchment. The result is likely to reflect the connectivity between static and lentic habitat and is not merely a question of the simple distance from the source populations. We also discuss how the potential distribution map can be applied to the management of piscivorous chub.     

Age,growth and habitat use of the topmouth gudgeon,Pseudorasbora parva in irrigation ditches on northwestern Kyushu Island,Japan

In Japan, Pseudorasbora parva has not been negatively affected by artificial changes in its habitat. Ecological traits of the species may explain this easy adaptation to artificial habitats. An ecological investigation of P. parva inhabiting irrigation ditches was conducted monthly for approximately two years on northwestern Kyushu Island, Japan. Measurements of 1927 individuals revealed two size classes, maturation age (1 year), and lifespan (1–2 years). Results of this study and previous studies show that the species exhibits life history plasticity. In addition, habitat use of ditches indicates that abundances correlated positively with water temperature and negatively with current velocity in the crop‐growing season, and positively with water depth and negatively with current velocity in the fallow season. Structural factors such as ditch revetment conditions and aquatic macrophyte cover were unimportant for this species. The study shows that the gudgeon adapts to artificial ditches easily and successfully because of both life history plasticity and wide‐scale use of the habitat.     

Non‐native salmonids affect amphibian occupancy at multiple spatial scales

Aim The introduction of non‐native species into aquatic environments has been linked with local extinctions and altered distributions of native species. We investigated the effect of non‐native salmonids on the occupancy of two native amphibians, the long‐toed salamander (Ambystoma macrodactylum) and Columbia spotted frog (Rana luteiventris), across three spatial scales: water bodies, small catchments and large catchments. Location Mountain lakes at ≥ 1500 m elevation were surveyed across the northern Rocky Mountains, USA. Methods We surveyed 2267 water bodies for amphibian occupancy (based on evidence of reproduction) and fish presence between 1986 and 2002 and modelled the probability of amphibian occupancy at each spatial scale in relation to habitat availability and quality and fish presence. Results After accounting for habitat features, we estimated that A. macrodactylum was 2.3 times more likely to breed in fishless water bodies than in water bodies with fish. Ambystoma macrodactylum also was more likely to occupy small catchments where none of the water bodies contained fish than in catchments where at least one water body contained fish. However, the probability of salamander occupancy in small catchments was also influenced by habitat availability (i.e. the number of water bodies within a catchment) and suitability of remaining fishless water bodies. We found no relationship between fish presence and salamander occupancy at the large‐catchment scale, probably because of increased habitat availability. In contrast to A. macrodactylum, we found no relationship between fish presence and R. luteiventris occupancy at any scale. Main conclusions Our results suggest that the negative effects of non‐native salmonids can extend beyond the boundaries of individual water bodies and increase A. macrodactylum extinction risk at landscape scales. We suspect that niche overlap between non‐native fish and A. macrodactylum at higher elevations in the northern Rocky Mountains may lead to extinction in catchments with limited suitable habitat.     

Complex interactions between native and invasive fish: the simultaneous effects of multiple negative interactions

We suggest that the ultimate outcome of interactions between native species and invasive species (extinction or coexistence) depends on the number of simultaneous negative interactions (competition and predation), which depends on relative body sizes of the species. Multiple simultaneous interactions may constrain the ability of native species to trade fitness components (i.e., reduced growth for reduced risk of predation) causing a spiral to extinction. We found evidence for five types of interactions between the adults and juveniles of introduced western mosquitofish (Gambusia affinis) and the juveniles of native least chub (Iotichthys phlegethontis). We added ten large (23–28 mm) and seven small (9–13 mm) young-of-the-year (YOY) least chub to replicate enclosures with zero, low, and high densities of mosquitofish in a desert spring ecosystem. Treatments with mosquitofish reduced the average survival of least chub by one-third. No small YOY least chub survived in enclosures with high mosquitofish densities. We also performed two laboratory experiments to determine mortality to predation, aggressiveness, and habitat selection of least chub in the presence of mosquitofish. Mean mortality of least chub due to predation by large mosquitofish was 69.7% over a 3-h trial. Least chub were less aggressive, selected protected habitats (Potamogeton spp.), and were more stationary in the presence of mosquitofish where the dominance hierarchy was large mosquitofish>>large least chubsmall mosquitofish>>small least chub. Least chub juveniles appear to be figuratively caught in a vice. Rapid growth to a size refuge could reduce the risk of predation, but the simultaneous effects of competition decreased least chub growth and prolonged the period when juveniles were vulnerable to mosquitofish predation.     

Ground-truthing the impact of invasive species: spatio-temporal overlap between native least chub and introduced western mosquitofish

Western mosquitofish (Gambusia affinis) are one of the most destructive introduced species in the American West. The negative impact of introduced species on native taxa depends on their spatio-temporal overlap, which will determine the availability of refugia for native species. Experiments on the mechanisms underlying the interactions between introduced and native species rarely address habitat use, overlap, and refugia because individuals are confined to enclosures. In a previous study we used cages, microcosms, and aquaria to show that mosquitofish could prey on and out-compete native least chub (Iotichthys phlegethontis). In this study, we examined the spatio-temporal overlap between mosquitofish and least chub under natural conditions. We found periods of overlap and partitioning in the seasonal and diel habitat use of these species. Both species used shallow habitats during the day and night throughout the spring when least chub were spawning. Predation by adult mosquitofish on young least chub during the spring likely explains the reduction in least chub recruitment in the presence of mosquitofish. During the summer least chub avoided mosquitofish by exclusively using cooler habitats, or by occupying deeper, cooler habitats during the day when mosquitofish were active, and shallower, warmer habitats at night when mosquitofish were inactive. A shift to cooler habitats in the presence of mosquitofish may result in decreased growth and fecundity of least chub. However, we suggest that a reduction of warm shallow habitat may have a disproportionately greater negative impact on mosquitofish than species native to temperate regions, such as least chub. Habitat manipulations that reduce or eliminate warm habitats may ameliorate the harmful effects of mosquitofish and promote the long-term persistence of native taxa.     

Irrigated urban trees exhibit greater functional trait plasticity compared to natural stands

Urbanization creates novel ecosystems comprised of species assemblages and environments with no natural analogue. Moreover, irrigation can alter plant function compared to non-irrigated systems. However, the capacity of irrigation to alter functional trait patterns across multiple species is unknown but may be important for the dynamics of urban ecosystems. We evaluated the hypothesis that urban irrigation influences plasticity in functional traits by measuring carbon-gain and water-use traits of 30 tree species planted in Southern California, USA spanning a coastal-to-desert gradient. Tree species respond to irrigation through increasing the carbon-gain trait relationship of leaf nitrogen per specific leaf area compared to their native habitat. Moreover, most species shift to a water-use strategy of greater water loss through stomata when planted in irrigated desert-like environments compared to coastal environments, implying that irrigated species capitalize on increased water availability to cool their leaves in extreme heat and high evaporative demand conditions. Therefore, irrigated urban environments increase the plasticity of trait responses compared to native ecosystems, allowing for novel response to climatic variation. Our results indicate that trees grown in water-resource-rich urban ecosystems can alter their functional traits plasticity beyond those measured in native ecosystems, which can lead to plant trait dynamics with no natural analogue.     

Bluehead chub nesting activity: a potential mechanism of population persistence in degraded stream habitats

Aquatic biodiversity continues to decline as humans modify the landscape, mainly because of stream habitat alterations and loss caused by urban development. Bluehead chubs may mitigate some effects of instream habitat degradation by providing clean gravel substrate via their spawning nests. We used path analysis, an extension of multiple linear regression, to explore the relationships among instream habitat degradation, adult chub abundance, chub nesting activity, and chub reproductive performance. Age-0 chub abundance was best explained by small adult abundance and nest abundance. Habitat disturbance indirectly and negatively influenced age-0 chub abundance through adult chubs and nest abundance. Percentages of pool and run habitat also had indirect negative effects on age-0 chub abundance. Several metrics of chub nesting activity (nest density [proportion of substrate occupied by nests], average nest size, and number of nests) were explained by both adult chub abundance and nesting site conditions. Variability among stream systems described significant variation in adult chub abundance and nesting characteristics and, if unaccounted for, would have resulted in large unexplained variability. Chub nesting activity served as a link between habitat degradation, adult chub abundance, and their reproductive performance. Our study provides preliminary evidence that bluehead chubs’ nesting activity may be a mechanism of their persistence in degraded stream reaches. We recommend confirmatory studies through in-stream manipulative experiments.     

Potential of restored gravel pits to provide suitable habitats for Eurasian otters in anthropogenic landscapes

In anthropogenic landscapes, which are usually characterized by the existence of highly heterogeneous patchworks of habitats with different conservation status, restored gravel pits have the potential to play a conservation role for semiaquatic species such as Eurasian otters (Lutra lutra). Here, we report an otter habitat suitability analysis on a complex fluvial system in an anthropogenic landscape to understand the role of different artificial water bodies (i.e. irrigation channels and ditches) and natural water bodies (i.e. rivers) related to a restored gravel pit lagoon system as providers of suitable habitats for otters. We implemented seven sampling campaigns during 2016 and 2017 consisting of 19 transects across all existing types of water bodies. We integrated 34 environmental variables with otter habitat use, measured by three spraint marking intensity indicators. We found that otter use of water bodies was not related to the natural or artificial origin of the water. Three key factors influenced habitat suitability: riparian vegetation tree cover, forestland use (either natural or planted) within 100 m from the banks, and the level of human disturbance. Our results suggest that otters' tolerance of human activities might be lower in key areas of their territory compared to areas they pass through or explore. We argue that restored gravel pit lagoon systems can potentially play a role in providing suitable habitats for otters in anthropogenic landscapes.     

Strong and parallel salinity‐induced phenotypic plasticity in one generation of threespine stickleback

Phenotypic plasticity is a major factor contributing to variation of organisms in nature, yet its evolutionary significance is insufficiently understood. One example system where plasticity might have played an important role in an adaptive radiation is the threespine stickleback (Gasterosteus aculeatus), a fish that has diversified after invading freshwater lakes repeatedly from the marine habitat. The parallel phenotypic changes that occurred in this radiation were extremely rapid. This study evaluates phenotypic plasticity in stickleback body shape in response to salinity in fish stemming from a wild freshwater population. Using a split‐clutch design, we detected surprisingly large phenotypically plastic changes in body shape after one generation. Fish raised in salt water developed shallower bodies and longer jaws, and these changes were consistent and parallel across families. Although this work highlights the effect of phenotypic plasticity, we also find indications that constraints may play a role in biasing the direction of possible phenotypic change. The slopes of the allometric relationship of individual linear traits did not change across treatments, indicating that plastic change does not affect the covariation of traits with overall size. We conclude that stickleback have a large capacity for plastic phenotypic change in response to salinity and that plasticity and evolutionary constraints have likely contributed to the phenotypic diversification of these fish.     

River alteration and niche overlap among three native minnows (Cyprinidae) in the Missouri River hydrosystem

The influence of physical and hydrologic stabilization on habitat niche overlap among three native cyprinid species: flathead chub Platygobio gracilis , sicklefin chub Macrhybopsis meeki and sturgeon chub Macrhybopsis gelida , in riverine segments of the Missouri and Yellowstone Rivers in western North Dakota and eastern Montana, was evaluated. Collectively the three species exhibited higher niche overlap in quasi‐natural river segments than in segments highly altered by a mainstem dam based on relatively high percentages of individuals in quasi‐natural river segments that were classified correctly, according to species, in discriminant function analyses of resource use, compared to lower percentages of individuals classified correctly in the altered river segments. The lower niche overlap in altered river segments resulted primarily from the lower overlap between flathead chub and the remaining species; this appears to be related to a decline in the diversity of natural habitats and conditions that provided a wide range of habitat conditions suitable for all three species. Results from this study suggest that selective segregation and habitat changes, rather than interactive segregation and competition, is probably the mechanism responsible for the pattern of habitat use and niche overlap among the three species in the altered segments.     

Urbanization drives contemporary evolution in stream fish

Human activities reduce biodiversity but may also drive diversification by modifying selection. Urbanization alters stream hydrology by increasing peak water velocities, which should in turn alter selection on the body morphology of aquatic species. Here, we show how urbanization can generate evolutionary divergence in the body morphology of two species of stream fish, western blacknose dace (Rhinichthys obtusus) and creek chub (Semotilus atromaculatus). We predicted that fish should evolve more streamlined body shapes within urbanized streams. We found that in urban streams, dace consistently exhibited more streamlined bodies while chub consistently showed deeper bodies. Comparing modern creek chub populations with historical museum collections spanning 50 years, we found that creek chub (1) rapidly became deeper bodied in streams that experienced increasing urbanization over time, (2) had already achieved deepened bodies 50 years ago in streams that were then already urban (and showed no additional deepening over time), and (3) remained relatively shallow bodied in streams that stayed rural over time. By raising creek chub from five populations under common conditions in the laboratory, we found that morphological differences largely reflected genetically based differences, not velocity–induced phenotypic plasticity. We suggest that urbanization can drive rapid, adaptive evolutionary responses to disturbance, and that these responses may vary unpredictably in different species.     

Morphological divergence between three Arctic charr morphs – the significance of the deep‐water environment

Morphological divergence was evident among three sympatric morphs of Arctic charr (Salvelinus alpinus (L.)) that are ecologically diverged along the shallow‐, deep‐water resource axis in a subarctic postglacial lake (Norway). The two deep‐water (profundal) spawning morphs, a benthivore (PB‐morph) and a piscivore (PP‐morph), have evolved under identical abiotic conditions with constant low light and temperature levels in their deep‐water habitat, and were morphologically most similar. However, they differed in important head traits (e.g., eye and mouth size) related to their different diet specializations. The small‐sized PB‐morph had a paedomorphic appearance with a blunt head shape, large eyes, and a deep body shape adapted to their profundal lifestyle feeding on submerged benthos from soft, deep‐water sediments. The PP‐morph had a robust head, large mouth with numerous teeth, and an elongated body shape strongly related to their piscivorous behavior. The littoral spawning omnivore morph (LO‐morph) predominantly utilizes the shallow benthic–pelagic habitat and food resources. Compared to the deep‐water morphs, the LO‐morph had smaller head relative to body size. The LO‐morph exhibited traits typical for both shallow‐water benthic feeding (e.g., large body depths and small eyes) and planktivorous feeding in the pelagic habitat (e.g., streamlined body shape and small mouth). The development of morphological differences within the same deep‐water habitat for the PB‐ and PP‐morphs highlights the potential of biotic factors and ecological interactions to promote further divergence in the evolution of polymorphism in a tentative incipient speciation process. The diversity of deep‐water charr in this study represents a novelty in the Arctic charr polymorphism as a truly deep‐water piscivore morph has to our knowledge not been described elsewhere.     

Kurze Notiz über Rivulus beniensis (Pisces,Cyprinodontidae) in seinem natürlichen Lebensraum bei Tingo Maria (ost‐andiner Hang in Peru)

Rivulus beniensis, formerly found by the author in small artificial ditches near Tingo Maria (Peru), is reproted from a natural habitat in the environment. The population in the ditches originates from this habitat. A chemical analysis of the water is included.     

Life-history traits of invasive fish in small Mediterranean streams

We compared the life-history traits of native and invasive fish species from Catalan streams in order to identify the characters of successful invasive fish species. Most of the exotic fish species were characterized by large size, long longevity, late maturity, high fecundity, few spawnings per year, and short reproductive span, whereas Iberian native species exhibited predominantly the opposite suite of traits. Species native to the southeastern Pyrenees watershed were also significantly different from species native to the rest of the Iberian Peninsula but not native to this watershed. Iberian exotic species come predominantly from large river basins, whereas Catalan streams (and other small, coastal river basins) correspond to basins and streams of a smaller size and different hydrology, with differences in species composition and life-history traits of fish. The occurrence and spread of invasive species was not significantly related to life-history traits but to introduction date. The successful prediction of future invasive species is limited due to small differences in life-history and ecological traits between native and exotic species. Fecundity, age at maturity, water quality flexibility, tolerance to pollution and habitat seem the most discriminating life-history variables. This revised version was published online in July 2006 with corrections to the Cover Date.     

Habitat patchiness affects distribution and microhabitat use of endangered Mira chub Squalius torgalensis (Actinopterygii,Cypriniformes)

We hypothesized that distribution and microhabitat use by imperilled chub Squalius torgalensis in the Torgal stream, Portugal, during low flows, were related to spatial patchiness in physical resources and shifts in ontogenetic preferences. We mapped fish abundance and sampled microhabitat use and availability via snorkelling. We used the coefficient of dispersion in abundance, and spatial autocorrelation analyses to characterize chub distribution, and Hurdle models to relate chub presence and abundance to habitat characteristics. We showed that chub displayed an aggregated distribution, apparently in association with patchily distributed and autocorrelated physical resources, such as debris, roots and aquatic vegetation. Microhabitat use generally was unrelated to velocity, and ontogenetic differences in microhabitat use were not substantial. Nevertheless, sometimes small chub preferred low-velocity patches with coarse substrata, debris and roots, whereas large chub preferred deeper patches with roots and aquatic vegetation. Results suggest that, in low flow conditions, chub respond to resource patchiness by congregating in favourable microhabitats, and that maintenance of mosaics of patches with variable substrata, cover and depth may be important for the persistence of fish in Mediterranean streams.     

Phenotypic plasticity of introduced versus native purple loosestrife: univariate and multivariate reaction norm approaches

The plastic responses to environmental change by Lythrum salicaria (purple loosestrife) were compared between native plants derived from seeds collected in Europe and those introduced into North America. Plants from nine populations each were grown under two levels of water and nutrient conditions. At the end of the growing season, samples were evaluated for eight traits related to their life history, plant size/architecture, and reproduction. Genetic (G), environmental (E), and G × E interactions were assessed by restricted maximum likelihood (REML) analysis of covariance (ANCOVA) and multivariate analysis of covariance (MANCOVA). Both univariate and multivariate reaction norm analyses were used to test for differences in the magnitude and direction of phenotypic plasticity between introduced and native plants. Under high-nutrient conditions, introduced plants were taller and had more branches and greater aboveground biomass. They also exhibited significantly greater amounts of phenotypic plasticity for aboveground biomass than did the natives in response to changing nutrient levels in standing water. This difference in univariate plasticity contributed to the general contrast in multivariate plasticity between introduced and native plants. These results support the idea that introduced plants may successfully invade a habitat and grow better than native plants in response to increased resources.     

Phenotypic plasticity of reproductive traits in response to food availability in invasive and native species of nematode

Invasive species cause severe ecological and economic damage; however, the mechanisms underlying their successful invasion often remain elusive. In the case of Bursaphelenchus xylophilus, a global quarantine pest which invaded Asia and Europe, it has been suggested that this species possesses highly competitive abilities, which promotes its establishment and rapid spread. To explore biological traits that may explain its highly competitive abilities, we focused on expression of phenotypic plasticity in response to the food conditions experienced by the females during their development as juveniles in the invasive species B. xylophilus and native species Bursaphelenchus mucronatus. We report an unexpected significant difference of phenotypic trade-off between egg number and egg size in the invasive species B. xylophilus and native species B. mucronatus. This leads to superior propagation ability of invasive species, under high and low food conditions in culture. These effects reflect adaptive optimal resource allocation where more eggs are produced in favorable environments to enhance population viability. Furthermore, we show that B. xylophilus eggs hatched earlier than B. mucronatus when their parents experienced high food availability. Thus, this study revealed, for the first time, phenotypic plasticity of reproductive traits in B. xylophilus which empowers the species a competitive advantage relative to their native counterpart B. mucronatus when they are under different range of food availability. These results are a step towards answering the vital question of how an exotic invasive species exclude a native species from its original niche.     

Adaptive divergence in Darwin's small ground finch (Geospiza fuliginosa): divergent selection along a cline

We examine here, in a single year (2005), phenotypic divergence along a 560‐m elevation gradient in Darwin's small ground finch (Geospiza fuliginosa) in the Galápagos Islands. In this sample, four composite measures of phenotypic traits showed significant differences along the 18‐km geographical cline extending from lowlands to highlands. Compared with lowland birds, highland birds had larger and more pointed beaks, and thicker tarsi, but smaller feet and claws. Finches in an intervening agricultural zone had predominantly intermediate trait values. In a second, mark–recapture study we analyse selection on morphological traits among birds recaptured across years (2000–2005) in lowland and highland habitats. Birds were more likely to survive in the highlands and during the wet season, as well as if they had large beaks and bodies. In addition, highland birds exhibited higher survival rates if they had small feet and pointed beaks – attributes common to highland birds as a whole. Lowland birds were more likely to survive if they possessed the opposite traits. Selection therefore reinforced existing morphological divergence, which appears to reflect local adaptation to differing resources during the predominantly drought‐ridden conditions that characterized the 5‐year study. Alternative explanations – including genetic drift, matching habitat choice, deformation by parasites, and the effects of wear – received little or no support. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 45–59.     

Adaptive divergence vs. environmental plasticity: tracing local genetic adaptation of metamorphosis traits in salamanders

In order to assess the significance of local adaptation relative to environmental plasticity on the evolution of life history traits, we analysed the possible genetic basis of differences between pond- and stream-breeding fire salamanders (Salamandra salamandra) in Germany. These salamanders typically deposit their larvae in small streams, where they grow until they are sufficiently large to metamorphose. However, some populations in Western Germany use ponds as larval habitat. Because habitat quality of streams differs from that of ponds one expects life history differences in the pond animals, which may result either from a plastic response or through genetic differentiation (i.e. local adaptation). Using a phylogeographical analysis of mitochondrial D-loop sequences, we show that both stream and pond populations in Western Germany are derived from a single lineage that recolonized following the last glaciation. This finding suggests that pond breeding originated very recently. Our studies of habitat quality and metamorphic behaviour of larvae in natural ponds and streams disclosed that pond larvae experience a significantly reduced food supply and greater risk of drying than do stream larvae. Pond larvae metamorphose earlier at the cost of reduced mass. Common-environment experiments with pond and stream larvae show that metamorphic behaviour of pond larvae under limited-food conditions is determined genetically and is not simply a plastic response to the differing habitat conditions. These results show that phenotypic plasticity is less important than local adaptation in explaining differences in ecological diversification within this species and suggests the possibility of rapid evolution of genetic adaptations when new habitats are exploited.