Winter survival of age-0 smallmouth bass, <Emphasis Type="Italic">Micropterus dolomieu</Emphasis>, in north eastern lakes

We examined the winter survival of age-0 smallmouth bass,Micropterus dolomieu, in lakes at their northern limit of distribution in New Brunswick, Canada. Pre- and post-winter collections over a 3 year period suggested the smallest individuals, <50 mm total length, died during winter under ice. Experiments with wild, age-0 smallmouth bass held in lakes demonstrated a size-dependent survival where smaller individuals had greater survival with an increasing temperature of exposure, but the largest individuals had reduced survival at warmer temperatures. Survivors lost 22–54% of pre-winter energy reserves measured as ash-free dry weight, which was similar to wild individuals captured in spring. Body size at the onset of winter affects survival, but it also appears that temperature interacting with a suite of environmental and biological factors affect changes in energy use and therefore survival of age-0 smallmouth bass under ice in winter.     

Quantifying the potential effects of climate change and the invasion of smallmouth bass on native lake trout populations across Canadian lakes

Climate change and invasive species are two stressors that should have large impacts on native species in aquatic and terrestrial ecosystems. We quantify and integrate the effects of climate change and the establishment of an invasive species (smallmouth bass Micropterus dolomieu ) on native lake trout Salvelinus namaycush populations. We assembled a dataset of almost 22 000 Canadian lakes that contained information on fish communities, lake morphologies, and geography. We examined the pelagic-benthic and littoral forage fish community available to lake trout populations across three lake size classes in these aquatic ecosystems. Due to the decreased presence of alternate prey resources, lake trout populations residing in smaller lakes are more vulnerable to the effects of smallmouth bass establishment. A detailed spatially and temporally explicit approach to assess smallmouth bass invasion risk in Ontario lakes suggests that the number of Ontario lakes with vulnerable lake trout populations could increase from 118 (~1%) to 1612 (~20%) by 2050 following projected climate warming. In addition, we identified nearly 9700 lake trout populations in Canada threatened by 2100, by the potential range expansion of smallmouth bass. Our study provides an integration of two major stressors of ecosystems, namely climate change and invasive species, by considering climate-change scenarios, dispersal rates of invasive species, and inter-specific biotic interactions.     

Spatial sorting drives morphological variation in the invasive bird, Acridotheris tristis

The speed of range expansion in many invasive species is often accelerating because individuals with stronger dispersal abilities are more likely to be found at the range front. This 'spatial sorting' of strong dispersers will drive the acceleration of range expansion. In this study, we test whether the process of spatial sorting is at work in an invasive bird population (Common myna, Acridotheris tristis) in South Africa. Specifically, we sampled individuals across its invasive range and compared morphometric measurements relevant and non-relevant to the dispersal ability. Besides testing for signals of spatial sorting, we further examined the effect of environmental factors on morphological variations. Our results showed that dispersal-relevant traits are significantly correlated with distance from the range core, with strong sexual dimorphism, indicative of sex-biased dispersal. Morphological variations were significant in wing and head traits of females, suggesting females as the primary dispersing sex. In contrast, traits not related to dispersal such as those associated with foraging showed no signs of spatial sorting but were significantly affected by environmental variables such as the vegetation and the intensity of urbanisation. When taken together, our results support the role of spatial sorting in facilitating the expansion of Common myna in South Africa despite its low propensity to disperse in the native range.     

Will northern fish populations be in hot water because of climate change?

Predicted increases in water temperature in response to climate change will have large implications for aquatic ecosystems, such as altering thermal habitat and potential range expansion of fish species. Warmwater fish species, such as smallmouth bass, Micropterus dolomieu , may have access to additional favourable thermal habitat under increased surface-water temperatures, thereby shifting the northern limit of the distribution of the species further north in Canada and potentially negatively impacting native fish communities. We assembled a database of summer surface-water temperatures for over 13 000 lakes across Canada. The database consists of lakes with a variety of physical, chemical and biological properties. We used general linear models to develop a nation-wide maximum lake surface-water temperature model. The model was extended to predict surface-water temperatures suitable to smallmouth bass and under climate-change scenarios. Air temperature, latitude, longitude and sampling time were good predictors of present-day maximum surface-water temperature. We predicted lake surface-water temperatures for July 2100 using three climate-change scenarios. Water temperatures were predicted to increase by as much as 18 °C by 2100, with the greatest increase in northern Canada. Lakes with maximum surface-water temperatures suitable for smallmouth bass populations were spatially identified. Under several climate-change scenarios, we were able to identify lakes that will contain suitable thermal habitat and, therefore, are vulnerable to invasion by smallmouth bass in 2100. This included lakes in the Arctic that were predicted to have suitable thermal habitat by 2100.     

Rapid evolution of larval life history,adult immune function and flight muscles in a poleward‐moving damselfly

Although a growing number of studies have documented the evolution of adult dispersal‐related traits at the range edge of poleward‐expanding species, we know little about evolutionary changes in immune function or traits expressed by nondispersing larvae. We investigated differentiation in larval (growth and development) and adult traits (immune function and flight‐related traits) between replicated core and edge populations of the poleward‐moving damselfly Coenagrion scitulum. These traits were measured on individuals reared in a common garden experiment at two different food levels, as allocation trade‐offs may be easier to detect under energy shortage. Edge individuals had a faster larval life history (growth and development rates), a higher adult immune function and a nearly significant higher relative flight muscle mass. Most of the differentiation between core and edge populations remained and edge populations had a higher relative flight muscle mass when corrected for latitude‐specific thermal regimes, and hence could likely be attributed to the range expansion process per se. We here for the first time document a higher immune function in individuals at the expansion front of a poleward‐expanding species and documented the rarely investigated evolution of faster life histories during range expansion. The rapid multivariate evolution in these ecological relevant traits between edge and core populations is expected to translate into changed ecological interactions and therefore has the potential to generate novel eco‐evolutionary dynamics at the expansion front.     

Predation of Japanese dace, Tribolodon hakonensis, by largemouth bass, Micropterus salmoides, in experimental aquaria

 To test the size range of prey fish that largemouth bass, Micropterus salmoides, can successfully consume, live Japanese dace, Tribolodon hakonensis, were given as prey fish to individual largemouth bass in aquaria. The ratio of maximum standard length (SL) of the Japanese dace consumed by largemouth bass was 46–69% of bass SL. The maximum length of Japanese dace consumed did not differ significantly between largemouth bass and smallmouth bass (M. dolomieu) previously studied, although largemouth bass have relatively larger mouth sizes than smallmouth bass. Largemouth bass occasionally injured and killed Japanese dace larger than the limit that could be consumed.     

Spatiotemporal patterns and habitat associations of smallmouth bass (Micropterus dolomieu) invading salmon‐rearing habitat

1. Smallmouth bass (Micropterus dolomieu) have been widely introduced to fresh waters throughout the world to promote recreational fishing opportunities. In the Pacific Northwest (U.S.A.), upstream range expansions of predatory bass, especially into subyearling salmon‐rearing grounds, are of increasing conservation concern, yet have received little scientific inquiry. Understanding the habitat characteristics that influence bass distribution and the timing and extent of bass and salmon overlap will facilitate the development of management strategies that mitigate potential ecological impacts of bass. 2. We employed a spatially continuous sampling design to determine the extent of bass and subyearling Chinook salmon (Oncorhynchus tshawytscha) sympatry in the North Fork John Day River (NFJDR), a free‐flowing river system in the Columbia River Basin that contains an upstream expanding population of non‐native bass. Extensive (i.e. 53 km) surveys were conducted over 2 years and during an early and late summer period of each year, because these seasons provide a strong contrast in the river’s water temperature and flow condition. Classification and regression trees were applied to determine the primary habitat correlates of bass abundance at reach and channel‐unit scales. 3. Our study revealed that bass seasonally occupy up to 22% of the length of the mainstem NFJDR where subyearling Chinook salmon occur, and the primary period of sympatry between these species was in the early summer and not during peak water temperatures in late summer. Where these species co‐occurred, bass occupied 60–76% of channel units used by subyearling Chinook salmon in the early summer and 28–46% of the channel units they occupied in the late summer. Because these rearing salmon were well below the gape limitation of bass, this overlap could result in either direct predation or sublethal effects of bass on subyearling Chinook salmon. The upstream extent of bass increased 10–23 km (2009 and 2010, respectively) as stream temperatures seasonally warmed, but subyearling Chinook salmon were also found farther upstream during this time. 4. Our multiscale analysis suggests that bass were selecting habitat based on antecedent thermal history at a broad scale, and if satisfactory temperature conditions were met, mesoscale habitat features (i.e. channel‐unit type and depth) played an additional role in determining bass abundance. The upstream extent of bass in the late summer corresponded to a high‐gradient geomorphic discontinuity in the NFJDR, which probably hindered further upstream movements of bass. The habitat determinants and upstream extent of bass were largely consistent across years, despite marked differences in the magnitude and timing of spring peak flows prior to bass spawning. 5. The overriding influence of water temperature on smallmouth bass distribution suggests that managers may be able limit future upstream range expansions of bass into salmon‐rearing habitat by concentrating on restoration activities that mitigate climate‐ or land‐use‐related stream warming. These management activities could be prioritised to capitalise on survival bottlenecks in the life history of bass and spatially focused on landscape knick points such as high‐gradient discontinuities to discourage further upstream movements of bass.     

Prey naivety in the behavioural responses of juvenile Chinook salmon (Oncorhynchus tshawytscha) to an invasive predator

1. Non‐native predators might inflict proportionally higher mortality on prey that have no previous experience of them, compared to species that have coexisted with the predator for some time. 2. We tested whether juvenile Chinook salmon (Oncorhynchus tshawytscha) were less able to recognise a non‐native than a native predator, by investigating behavioural responses to the chemical cues of the invasive smallmouth bass (Micropterus dolomieu) and the native northern pikeminnow (Ptychocheilus oregonensis) in both laboratory and field experiments. 3. Laboratory results demonstrated strong innate antipredator responses of individual juvenile Chinook salmon to northern pikeminnow; fish spent 70% of time motionless and exhibited 100% greater panic response than in controls. By contrast, antipredator responses to the chemical cues of smallmouth bass did not differ from controls. 4. These results were supported by similar differences in recognition of these predator odours by groups of juvenile Chinook salmon in fully natural conditions, though responses reflected a greater range of antipredator behaviours by individuals. In field trials, responses to northern pikeminnow odour resulted in increased flight or absence, reductions in swimming and foraging, and increased time spent near the substratum, compared to smallmouth bass odour. 5. Given that survival of juvenile fish is facilitated by predator recognition, our results support the hypothesis that naivety may be an important factor determining the effect of non‐native predators on prey populations. Efforts to manage the effect of native and non‐native predators may benefit by considering complex behavioural interactions, such as these at the individual and group levels.     

Fine scale spatio-temporal life history shifts in an invasive species at its expansion front

The dynamic shifts in life history traits by non-native populations at an invasion front may be driven by a combination of phenotypic plasticity and micro-evolutionary processes, allowing for appropriate responses to changes in biotic and abiotic factors associated with range expansion. The temporal and spatial scale at which these shifts occur, however, is largely unexplored. We examined the age-specific life-history responses of female round goby (Neogobius melanostomus) across a fine spatial scale by comparing traits of individuals at the expanding front of an invasive pathway with those of individuals living at the previous year’s front. Females in the new front were younger, exhibited higher body condition index and fecundity, and greater reproductive allocation than those living at the previous year’s front. No difference in back-calculated length at age or in seasonal growth increment was found between occupants of the old and new fronts. Age-specific differences in reproductive investment were observed; age-1 females from the new front exhibited greater reproductive effort but similar reproductive allocation, whereas age-2 females showed greater reproductive allocation in the new front, but similar reproductive effort. These age-specific differences may be related to differences in population density and other biotic factors encountered during ontogeny, or to micro-evolutionary processes. Our results demonstrate that fine scale changes in energy allocation towards reproduction through increases in reproductive allocation and fecundity occur at the invasion front, highlight how quickly those shifts can occur at the expansion front, and suggest that such changes in newly colonised areas facilitate range expansion.     

Expanding population edges: theories,traits, and trade‐offs

Recent patterns of global change have highlighted the importance of understanding the dynamics and mechanisms of species range shifts and expansions. Unique demographic features, spatial processes, and selective pressures can result in the accumulation and evolution of distinctive phenotypic traits at the leading edges of expansions. We review the characteristics of expanding range margins and highlight possible mechanisms for the appearance of phenotypic differences between individuals at the leading edge and core of the range. The development of life history traits that increase dispersal or reproductive ability is predicted by theory and supported with extensive empirical evidence. Many examples of rapid phenotypic change are associated with trade‐offs that may influence the persistence of the trait once expansion ends. Accounting for the effects of edge phenotypes and related trade‐offs could be critical for predicting the spread of invasive species and population responses to climate change.     

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

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Complete mitochondrial DNA sequences of two haplotypes of the smallmouth bass, <Emphasis Type="Italic">Micropterus dolomieu</Emphasis>, collected from nonindigenous populations in Japan

We determined the complete mitochondrial genome sequences of two haplotypes of the smallmouth bass, Micropterus dolomieu; individuals used in the analysis were collected from nonindigenous populations in Japan. Both genomes comprised 16,488 bp, with genome contents and gene orders being identical to those of other teleost fishes. A previous study revealed that the Japanese smallmouth bass had only two haplotypes, and the present study revealed that the complete mitochondrial DNA (mtDNA) sequences of the haplotypes differed in only one nucleotide difference. The low genetic diversity in the mtDNA of the smallmouth bass individuals in our study and the results of the mtDNA sequence comparison between the Japanese and the North American individuals suggested that the fish had been transplanted from a fish farm with a low-diversity stock.     

Precocious reproduction increases at the leading edge of a mangrove range expansion

Climate change‐driven shifts in species ranges are ongoing and expected to increase. However, life‐history traits may interact with climate to influence species ranges, potentially accelerating or slowing range shifts in response to climate change. Tropical mangroves have expanded their ranges poleward in the last three decades. Here, we report on a shift at the range edge in life‐history traits related to reproduction and dispersal. With a common garden experiment and field observations, we show that Rhizophora mangle individuals from northern populations reproduce at a younger age than those from southern populations. In a common garden at the northern range limit, 38% of individuals from the northernmost population were reproductive by age 2, but less than 10% of individuals from the southernmost population were reproductive by the same age, with intermediate amounts of reproduction from intermediate latitudes. Field observations show a similar pattern of younger reproductive individuals toward the northern range limit. We also demonstrate a shift toward larger propagule size in populations at the leading range edge, which may aid seedling growth. The substantial increase in precocious reproduction at the leading edge of the R. mangle range could accelerate population growth and hasten the expansion of mangroves into salt marshes.     

Does shelter influence the metabolic traits of a teleost fish?

Availability of shelter is an important component of habitat selection for animals as it can influence survival (protection against harsh physical conditions and predation) and growth (energy acquisition and expenditure). Few studies address the effect of shelter on metabolic expenditures associated with non-mechanical tasks (excluding station holding or movement). The main goal of this study was to investigate the influence of shelter use on metabolic traits of smallmouth bass (Micropterus dolomieu) from two populations (Kiamika River and Lake Long). Respirometry experiments on smallmouth bass were conducted to measure standard metabolic rate (SMR), resting metabolic rate (RMR), aerobic scope (AS), recovery time (RT) and excess post-exercise oxygen consumption (EPOC) in the presence or absence of shelter. The presence of shelter did not affect most metabolic traits, except for RMR, which was reduced in the presence of shelter for Lake Long fish. The results of this study also show that larger fish had lower SMR in the presence of shelter than when it was absent. When accounting for social hierarchy, there were no differences in most metabolic traits in dominant or subordinate fish in the presence or absence of shelter, except for RT, which was significantly lower in the presence of shelter for dominant fish. These results do not support the existence of an unequivocal relationship between individual metabolic traits and the presence of shelter. If physiological motives may influence the use of shelter, sheltering in itself might not have important consequences on energy expenditures required for non-mechanical tasks.     

What factors shape female phenotypes of a poleward‐moving damselfly at the edge of its range?

Individuals at the expansion front during a climate‐driven range expansion are expected to differ phenotypically from those individuals in core populations. Little information is known about the joint, potentially opposing, effects of stressful conditions at the range edge versus evolutionary changes that take place during range expansion in shaping the phenotypes at the range front. We investigated the effect of range expansion on immune function, body condition and flight‐related morphology (flight muscle ratio, wing loading, and wing aspect ratio) of field‐collected females of the poleward‐moving damselfly Coenagrion scitulum. Individuals at the expansion front had a lower body condition, which indicated more stressful conditions at the range edge. Despite the counteracting effect of the shorter growth season, the higher flight muscle ratios at the expansion front indicated a strong selection for dispersal ability during range expansion. The current study suggests that models need to incorporate the interplay of stressful conditions and evolutionary processes at the expansion front to arrive at robust predictions of future species distributions under global warming. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 556–568.     

Empirical assessment of potential interactions between walleye and smallmouth bass

The expansion of smallmouth bass Micropterus dolomieu throughout North America has created substantial changes in some food webs and created novel interactions with native predators in many systems through competition and predation, causing declines in native predator populations. In this study, we examined potential interactions between smallmouth bass (introduced) and walleye Sander vitreus (a native predator) in two Missouri River reservoirs and four natural lakes in South Dakota, USA, using historical standardized sampling data, which indexed four fitness‐related factors (relative abundance, size structure, condition, and growth) of both species. Of the 24 total relationships tested among all water bodies, three (13%) were determined to be significant (size structure in Pickerel Lake: P = 0.06; condition in Lake Kampeska: P = 0.06, and growth at Brant Lake: P = 0.02). Of these three, two relationships were negative as would be expected if negative interactions exist between these two species (size structure in Pickerel Lake: r = ?0.61; condition in Lake Kampeska: r = ?0.61); the remaining relationship was positive (growth at Brant Lake: r = 0.75). Further, no water body consistently demonstrated significant negative interactions in all relationships tested. The results of our study did not support the concept that introduced smallmouth bass interfere with walleye populations in South Dakota waters. Rather, it is more likely that other environmental factors influence the observed variability in walleye demographics and dynamics observed in this study. At this time, we believe that both fisheries will continue to co‐exist. However, continuous monitoring of both species is warranted as changing environmental conditions in the future could allow these interspecific relationships to shift toward the dominance of smallmouth bass over walleye.     

Spawning and Brood Defense of Smallmouth Bass Under the Process of Invasion into a Novel Habitat

Smallmouth bass, Micropterus dolomieu, known as one of the world's most disastrous invasive species, was introduced into Lake Aoki, central Japan in the last 10 years and is of concern for the conservation of the native biodiversity. We investigated spawning and brood defense under novel conditions to devise measures to eradicate a local population of this species. Compared to their native habitat, the size distribution of nesting males was less skewed to the right in this invasive population but rather a skewed normal distribution probably due to prevention of younger cohorts entering the nesting stock. The number of offspring deposited in nests was not positively related to the size of the nesting male. These observations imply that a shortage of suitable nest sites leads to an escalation in male–male competition, followed by a relaxation of female mate choice based on male size. Further, when presented with a model of a potential predator of the offspring, nesting males reacted individually and the intensity of site tenacity was independent of the male size. We propose that the removal of cover accompanied by the use of native predators of young bass can effectively decelerate the expansion of invasive smallmouth bass.     

Seasonal pattern of depth selection in smallmouth bass

The current study used a stationary acoustic telemetry array to monitor the depth selection of adult smallmouth bass Micropterus dolomieu in a large, oligotrophic lake in Ontario, Canada. At an annual scale, smallmouth bass demonstrated regular, seasonal changes in inhabited depths: fish occupied shallow water during the summer (2–5 m depth) and descended to deeper water (12–15 m depth) during winter under ice. Smallmouth bass remained above the thermocline in the summer, seasonal depth patterns did not vary across fish size and movements to and from seasonal depths were closely linked to the development and degradation of the thermocline. At finer time scales, smallmouth bass exhibited diel vertical migrations in summer, with fish moving to <2 m at night, and then descending to 3–5 m during the day. This pattern remained constant during the summer period examined, and varied with size such that larger fish remained deeper than smaller fish. During winter, depth did not vary across the 3-month monitoring period ( c . 14 m), but small (<2 m) changes in depth were observed periodically, suggesting limited movements were occurring. Results are further discussed in the context of climate change and reproductive success for this species.     

The role of progeny quality and male size in the nesting success of smallmouth bass: integrating field and laboratory studies

Smallmouth bass display size-specific variation in reproductive success with larger brood-guarding males in a population more likely to rear offspring to independence than smaller individuals. The exact mechanisms responsible for this size-specific increase in reproductive output have yet to be identified. To assist in this process, we investigated the relationship between the size of brood-guarding male smallmouth bass and offspring quality (in this case, egg physiology, egg morphology, egg size, hatching success and lab survival). Further, we examined how factors such as egg physiology, egg morphology and egg size influenced reproductive success in the wild and hatching success in a controlled laboratory environment. Nesting male smallmouth bass that successfully reared their offspring to independence spawned earliest in the nesting period were the largest individuals, and guarded eggs with greater concentrations of cortisol compared to males that abandoned their offspring prematurely. Offspring survival in the laboratory was not correlated with offspring survival in the wild, indicating that caution should be used interpreting studies that attempt to relate laboratory-derived survival metrics to the wild. Together, results demonstrate size-specific differences in offspring quality for nesting smallmouth bass, which are correlated with higher concentrations of cortisol in eggs. However, hatching success under laboratory conditions was dissimilar to nesting success in the field relative to cortisol concentrations.     

Description of two new gill myxozoans from smallmouth (Micropterus dolomieu) and largemouth (Micropterus salmoides) bass

Two previously undescribed species of myxozoan parasites were observed in the gills of bass inhabiting the Potomac and James River basins. They are described using morphological characteristics and small-subunit (SSU) rDNA gene sequences. Both were taxonomically identified as new species of Myxobolus; Myxobolus branchiarum n. sp. was found exclusively in smallmouth bass, and Myxobolus micropterii n. sp. was found in largemouth and smallmouth bass. Small, spherical, white plasmodia of M. branchiarum from smallmouth bass were observed grossly in the gills; these plasmodia had an average length of 320.3 μm and width of 246.1 μm. The development of the plasmodia is intralamellar in the secondary lamellae of the gills. Mature spores were pyriform in shape with a length of 12.8 ± 1.4 (8.1-15.1) μm and width of 6.9 ± 1.1 (4.0-9.0) μm. Analysis of SSU rDNA identified M. branchiarum in a sister-group to 3 species of Henneguya , although morphologically caudal appendages were absent. Myxobolus micropterii observed in the gills of largemouth and smallmouth bass had larger, ovoid, cream-colored plasmodia with an average length of 568.1 μm and width of 148.1 μm. The cysts developed at the distal end of the gill filament within the primary lamellae. The mature spores were ovoid in shape with a length of 10.8 ± 0.7 (9.2-12.2) μm and width of 10.6 ± 0.6 (9.0-11.8) μm. SSU rDNA analysis placed M. micropterii in a sister group with Henneguya lobosa and Myxobolus oliveirai . The highest prevalence of M. branchiarum was observed in the gills of bass collected from the Cowpasture River (50.9%). Prevalence was 44.6% in bass from the Potomac River and only 4.3% in bass collected from the Shenandoah River. A seasonal study of M. branchiarum , which included both infected and uninfected smallmouth bass, determined that a significantly higher intensity was observed in the spring than in the summer (P < 0.001) or fall (P = 0.004). In an analysis excluding uninfected bass, a higher intensity was observed in the spring than in the summer (P = 0.001) or fall (P = 0.008). Prevalence and seasonal differences were not determined for M. micropterii .