Out of the pot and into the fire: Explaining the vulnerability of an endangered small headwater stream fish to black‐bass Micropterus spp. invasion

Introduced predatory fishes have had consistently severe consequences for native fishes in stream environments around the world, although the drivers of these effects are often unclear. In the Swartkops River headwaters in South Africa, native Eastern Cape redfin Pseudobarbus afer were always absent from sites occupied by non‐native black basses Micropterus salmoides and Micropterus dolomieu, but generally co‐occurred with the native predators Anguilla marmorata and Anguilla mossambica. A natural experiment provided by flood‐mediated recolonization of black‐bass occupied sites by P. afer demonstrated depletion in black‐bass invaded sites. Field behavioural observations of P. afer indicated that they foraged among benthic cover during the day, but suspended in open water at night. As the nocturnal A. marmorata and A. mossambica foraged actively within structural cover at night and M. dolomieu and M. salmoides are diurnal or crepuscular predators, P .afer is thus optimized to avoid predation by native anguillid predators and not the functionally unique predatory black basses. The integration of distributional, temporal population dynamics and behavioural data suggests that the severe effects of Micropterus spp. are probably a consequence of prey naïveté and behaviour evolved to evade native predators.     

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.     

Invasion status of Florida bass Micropterus floridanus (Lesueur, 1822) in South Africa

Largemouth bass Micropterus salmoides are a popular North American angling species that was introduced into South Africa in 1928. To enhance the largemouth bass fisheries, Florida bass Micropterus floridanus were introduced into KwaZulu Natal, South Africa, in 1980. Knowledge on the status of M. floridanus in South Africa is required, because it lives longer and reaches larger sizes than M. salmoides, which may result in heightened impacts on native biota. Because M. floridanus are morphologically similar, but genetically distinct from M. salmoides, the distribution of this species was assessed by genetically screening 185 Micropterus sp. individuals sampled from 20 localities across South Africa using the mitochondrial ND2 gene. Individuals with mitochondrial DNA matching M. salmoides were recovered from 16 localities, whereas M. floridanus mitochondrial DNA was recovered from 13 localities. At nine localities (45%), the mitochondrial DNA of both species was detected. These results demonstrate M. floridanus dispersal to multiple sites across South Africa.     

Identification of centrarchid hepcidins and evidence that 17β-estradiol disrupts constitutive expression of hepcidin-1 and inducible expression of hepcidin-2 in largemouth bass (Micropterus salmoides)

Hepcidin is a highly conserved antimicrobial peptide and iron-regulatory hormone. Here, we identify two hepcidin genes (hep-1 and hep-2) in largemouth bass (Micropterus salmoides) and smallmouth bass (Micropterus dolomieu). Hepcidin-1 contains a putative ATCUN metal-binding site in the amino-terminus that is missing in hepcidin-2, suggesting that hepcidin-1 may function as an iron-regulatory hormone. Both hepcidins are predominately expressed in the liver of largemouth bass, similar to other fish and mammals. Experimental exposure of pond-raised largemouth bass to 17β-estradiol and/or the bacteria Edwardsiella ictaluri led to distinct changes in expression of hep-1 and hep-2. Estradiol reduced the constitutive expression of hep-1 in the liver. Bacterial exposure induced expression of hep-2, suggesting that hepcidin-2 may have an antimicrobial function, and this induction was abolished by estradiol. To our knowledge, this is the first report of the regulation of hepcidin expression by estradiol in either fish or mammals.     

The predatory impact of invasive alien smallmouth bass,Micropterus dolomieu (Teleostei: Centrarchidae), on indigenous fishes in a Cape Floristic Region mountain stream

Fish populations in the Rondegat River, a mountain stream in the Olifants-Doring system in the Cape Floristic Region, South Africa were surveyed to assess the impact of predatory alien invasive smallmouth bass Micropterus dolomieu (Lacepède) on the indigenous fishes. This was the first such attempt to quantify the predatory impacts of M. dolomieu within this region. The Rondegat River is home to five species of indigenous fish and is partially invaded by M. dolomieu, which has penetrated the lower river up to a waterfall barrier. Seasonal surveys were conducted at five sites above, and five below, the waterfall. Physical habitat was measured at each site. Four of the five indigenous fish species were absent at bass-invaded sites. Labeobarbus capensis (Smith), while still present below the waterfall, appeared to have suffered a near-total loss of post-spawning recruits. Analyses of physical habitat quality failed to explain the loss of indigenous fish below the waterfall, although sedimentation may have increased the vulnerability of the catfish Austroglanis gilli (Barnard) to M. dolomieu predation by obliterating benthic cover. Consequently, predation by M. dolomieu was presumed to be the critical mechanism explaining the loss of indigenous fishes in the lower Rondegat River.     

Species-specific impact of introduced largemouth bass Micropterus salmoides in the Groot Marico Freshwater Ecosystem Priority Area,South Africa

Largemouth bass Micropterus salmoides are among the world's 100 worst invaders and negatively affect aquatic biodiversity in many regions worldwide. In South Africa there is a paucity of empirical studies describing their impacts. The impact of M. salmoides on the fish community in the Groot Marico River catchment, an otherwise near-pristine river ecosystem and a freshwater ecosystem priority area, was assessed from surveys conducted in 2012. Fish presence and abundance were enumerated using multiple survey techniques, and their association with key habitat variables and the presence or absence of M. salmoides were assessed. A total of 14 native fish species were recorded, besides introduced M. salmoides, which occupied the majority of the mainstem and several tributaries downstream of barriers to upstream movement. Canonical correspondence analysis showed that only one native species, the Marico barb Barbus motebensis, had a negative spatial association with M. salmoides. Assessment of relative distributions showed this species to be excluded from M. salmoides-invaded river reaches, whereas the other native species were not visibly affected by the invader. This species-specificity of the impact of M. salmoides indicates that their impacts in South African streams may be dependent on predator-naiveté of prey.     

The effect of largemouth bass Micropterus salmoides on aquatic macro-invertebrate communities in the Wit River,Eastern Cape,South Africa

Fish predation is one of the driving forces of freshwater invertebrate community structures, with alien predators having a pronounced effect. A quantitative assessment of aquatic invertebrates in the Wit River, Sundays River catchment, Eastern Cape, South Africa, was undertaken to assess the impact of the alien fish Micropterus salmoides on their relative abundance and community structure. Communities in the stones-in-current and marginal vegetation biotopes were sampled in the presence and absence of M. salmoides in late summer (February–April) 2008. Results suggest that the presence of M. salmoides does have an impact on indigenous macroinvertebrate fauna and communities. Community structure in the stones-in-current biotopes did not differ significantly between sections of the river with or without fish. However, there was a significant difference in community structure in marginal vegetation between sections of river. In sections with M. salmoides several large or conspicuous taxa (Odonata, Hemiptera and Coleoptera) were significantly reduced (p < 0.05) or even absent, while cryptic/inconspicuous taxa (Trichoptera, Leptoceridae and Mollusca, Physidae) were significantly more abundant (p < 0.05).     

Life‐history variability of non‐native centrarchids in regulated river systems of the lower River Guadiana drainage (south‐west Iberian Peninsula)

Life‐history variability of two non‐native centrarchids, pumpkinseed Lepomis gibbosus and largemouth bass Micropterus salmoides, was evaluated in three stream stretches of the lower River Guadiana drainage (south‐west Iberian Peninsula) with different degrees of regulated flows. Abundance, condition and population structure differed among populations for both species, but invasion success was lower in the least regulated river. Lepomis gibbosus were abundant and had multiple age classes in the three river sites, whereas M. salmoides were less abundant and mainly represented by young‐of‐the‐year fish. Juvenile growth in L. gibbosus was similar in all three populations, though longevity was slightly greater in the population from the River Guadiana mainstream. Lepomis gibbosus exhibited a long reproductive season, but the duration of season, size at maturity and reproductive effort varied among populations. The life‐history differences found demonstrate the importance of species adaptation to local conditions which might favour their invasion success. Lepomis gibbosus were more adaptable and resilient to local conditions, whereas M. salmoides seemed dependent on reservoirs and large rivers for maintenance of riverine populations.     

Correlations between body length and otolith size in smallmouth bass Micropterus dolomieu Lacépède, 1802 with implications for retrospective growth analyses

Reconstructing individual growth history from analysis of increments in otoliths, scales, or spines can provide information on past growth responses to environmental variation, which in turn can be useful for predicting population‐level response to climate change. The objective of this study was to examine correlations between body length and different metrics of otolith size for Micropterus dolomieu. Three metrics corresponding to commonly‐used microstructural and ultrastructural otolith dimensions were measured using image analysis of digital micrographs from a sample of 214 M. dolomieu ranging from 115 to 438 mm total length collected in 2011–2013. It was found that anteroposterior length of whole otoliths provided much improved regression relationships with body size as well as ease of data collection and faster sampling throughput compared with microstructural measures from polished sections. When applying these metrics to reconstruct growth history the biological intercept model generally produced more reasonable back‐calculated estimates of length‐at‐age, although this was not consistent across all otolith metrics. Results suggest that whole otolith measures should be employed due to efficiency of data collection and greater reliability for reconstructing growth history in M. dolomieu.     

Complementarity of thermoregulatory rhythms in Micropterus salmoides and M. dolomieui

Largemouth (Micropterus salmoide) and smallmouth (M. dolomieui) blackbasses tested in an electronic shuttlebox exhibited behavioral thermoregulatory rhythms which were temporally complementary. With a LD 12 : 12 photoperiod, M. dolomieui exhibited a preferred-temperature peak of 30.1°C during the latter portion of the photophase, when M. salmoides reached a minimum of 27.1°C. M. dolomieui exhibited a minimum of 26.6°C during the latter portion of scotophase, while M. salmoides remained at a significantly higher plateau of about 29°C, with a peak of 29.5°C at the midpoint of scotophase. The phase relations of the thermoregulatory rhythms relative to photoperiod suggest that they are endogenously timed circadian rhythms entrained by photoperiod. The thermotemporal complementarity of these rhythms suggests an aspect of niche segregation between these largely sympatric congeneric species.     

The abiotic and biotic factors limiting establishment of predatory fishes at their expanding northern range boundaries in Ontario,Canada

There is a poor understanding of the importance of biotic interactions in determining species distributions with climate change. Theory from invasion biology suggests that the success of species introductions outside of their historical ranges may be either positively (biotic acceptance) or negatively (biotic resistance) related to native biodiversity. Using data on fish community composition from two survey periods separated by approximately 28 years during which climate was warming, we examined the factors influencing the establishment of three predatory centrarchids: Smallmouth Bass (Micropterus dolomieu), Largemouth Bass (M. salmoides), and Rock Bass (Ambloplites rupestris) in lakes at their expanding northern range boundaries in Ontario. Variance partitioning demonstrated that, at a regional scale, abiotic factors play a stronger role in determining the establishment of these species than biotic factors. Pairing lakes within watersheds where each species had established with lakes sharing similar abiotic conditions where the species had not established revealed both positive and negative relationships between the establishment of centrarchids and the historical presence of other predatory species. The establishment of these species near their northern range boundaries is primarily determined by abiotic factors at a regional scale; however, biotic factors become important at the lake‐to‐lake scale. Studies of exotic species invasions have previously highlighted how spatial scale mediates the importance of abiotic vs. biotic factors on species establishment. Our study demonstrates how concepts from invasion biology can inform our understanding of the factors controlling species distributions with changing climate.     

Age,growth and reproduction of non‐native largemouth bass Micropterus salmoides (Lacépède, 1802) populations in two temperate African impoundments

This study aims to improve the understanding of the establishment success of Micropterus salmoides by assessing growth, maturity and reproductive seasonality of two populations from the temperate region in South Africa. Results are compared to those from other populations to test previously proposed hypotheses that the (i) extent of the spawning season, (ii) longevity, and (iii) growth performance are environmentally temperature dependent. Micropterus salmoides were sampled monthly between March 2010 and March 2012 from Wriggleswade and Mankazana impoundments using either angling or multi‐meshed (35, 45, 47, 57, 65, 73, 77, 93, 106, 118, 150 and 153 mm) multifilament gillnet fleets, and a 1 m seine net. The assessed populations in South Africa performed as expected compared to other introduced populations. Age at maturity (1.3–2 years) and length at maturity (232–254 mm FL) were similar to other temperate populations. A well‐defined, spring spawning season conformed to findings in other temperate populations. Using sectioned sagittal otoliths it was shown that fish were long‐lived (14 years). Growth rates differed significantly between the two assessed populations but both were slower growing than tropical and subtropical introduced populations. Growth performance (Φ′: 2.77–2.99) was similar to other temperate populations. These results indicate that the biology of introduced M. salmoides populations in South Africa is comparable to those of other temperate populations and supports the environmental temperature‐dependent hypothesis.     

Influence of largemouth bass,Micropterus salmoides,on abundance and habitat selection of Cape galaxias,Galaxias zebratus,in a mountain stream in the Cape Floristic Region,South Africa

Predatory alien fishes have been widely introduced into streams in the Cape Floristic Region (CFR), South Africa, but little is known about their effect on native fishes. Results from this 2006 study suggest that the presence of alien predatory largemouth bass, Micropterus salmoides, may have influenced abundance and habitat selection of the native Galaxias zebratus at one location in a small CFR mountain stream. Numbers of adults, but not of juveniles, were significantly lower where M. salmoides was present, suggesting a size-specific influence on G. zebratus abundance. Because other studies have found the influence of a predator to be affected by prey size and the diel activity of predator and prey, we measured microhabitat use by adult and juvenile G. zebratus by day and by night. Galaxias zebratus selected deeper, faster-flowing microhabitats more strongly where M. salmoides was present than where it was absent. This suggests that G. zebratus adjusts its habitat use in the presence of M. salmoides, although differences in available habitat, or in interactions with other indigenous species, could be partly responsible for the observed differences. In-stream vegetation presence was strongly positively related to depth and strongly negatively related to velocity at positions where G. zebratus was observed, suggesting that the deeper, slower-flowing microhabitats occupied by G. zebratus were structurally more complex than the shallower, faster-flowing ones.     

Otolith chemistry of prey fish consumed by a fish predator: does digestion hinder Russian doll techniques?

The effect of digestion by a predatory fish (largemouth bass Micropterus salmoides) on stable isotopic (δ¹³C and δ¹⁸O) and trace elemental (Sr:Ca and Ba:Ca) compositions of prey fish (bluegill Lepomis macrochirus) otoliths was investigated in a laboratory experiment. Trace element and stable‐isotopic signatures of L. macrochirus otoliths were not significantly altered for up to 16 h after L. macrochirus were consumed by M. salmoides. Prey fish otoliths recovered from predator digesta can retain environmental stable isotopic and trace elemental signatures, suggesting that determination of environmental history for prey fishes by stable‐isotope and trace‐element analysis of otoliths recovered from stomachs of piscivorous fishes will be feasible.     

Utilizing stomach content and faecal DNA analysis techniques to assess the feeding behaviour of largemouth bass Micropterus salmoides and bluegill Lepomis macrochirus

In this study, the feeding behaviour of the non‐native invasive predatory fishes largemouth bass Micropterus salmoides and bluegill Lepomis macrochirus was studied in the Ezura River, a northern tributary of Lake Biwa, Japan. Prey composition was estimated based on visual examination of stomach contents and faecal DNA analysis to determine feeding habits of these predatory fishes. Stomach content analysis showed that native fishes (e.g. ayu Plecoglossus altivelis and gobies Rhinogobius spp.) and shrimps (e.g. Palaemon paucidens) were the major prey items for M. salmoides, while snails, larval Chironomidae and submerged macrophytes were the dominant prey items of L. macrochirus. Micropterus salmoides tended to select larger fish in the case of crucian carp Carassius spp., but smaller fishes in the case of P. altivelis and Rhinogobius spp. Faecal DNA analyses revealed prey compositions similar to those identified in predator stomach contents, and identified additional prey species not detected in stomach content inspection. This study demonstrated that both stomach content inspection and DNA‐based analysis bear several inherent shortcomings and advantages. The former method is straightforward, although identification of species can be inaccurate or impossible, whereas the latter method allows for accurate species identification, but cannot distinguish prey size or stage. Hence, integration of morphology‐based and DNA‐based methods can provide more reliable estimates of foraging habits of predatory fishes.     

A review of length–weight relationships of fish from the Segura River basin (SE Iberian Peninsula)

We present the relationships between fork length and total weight for 14 fish species from the Segura River basin (southeastern Spain): Barbus sclateri, Chondrostoma polylepis, Gobio lozanoi, Aphanius iberus, Micropterus salmoides, Lepomis gibbosus, Carassius auratus auratus, Cyprinus carpio carpio, Sander lucioperca, Gambusia holbrooki, Liza ramado, Mugil cephalus, Atherina boyeri and Pomatoschistus marmoratus. Significant length–weight relationships were found for all species. The values of the exponent b of the length–weight relationships ranged from 3.82 for Sander lucioperca to 2.59 for Micropterus salmoides.     

Variation in Prey-resource Utilization and Oral Jaw Gape Between two Populations of Largemouth Bass, Micropterus Salmoides

Ontogenetic changes in diet and jaw gape were compared between two indigenous populations of largemouth bass, Micropterus salmoides, to test the hypothesis that ecomorphology varies among broadly distributed fish populations. Two hundred seventy-two temperate (southwestern Michigan) M. salmoides and 265 subtropical (east-central Florida) conspecifics were analyzed for food habits and oral jaw gape height and width. Percent volumetric contributions of four functional prey categories (plankton, insect, crustacean, and fish) were compared among fish-size classes to determine if interval-specific differences in prey consumption existed between populations. Subtropical M. salmoides shifted from feeding on plankton and macroinvertebrates to fish by 20mm standard length (SL); and stopped consuming plankton by 29mm SL. Temperate largemouth bass did not become piscivorous until 37mm SL and continued utilizing plankton up to 69mm SL. Following the onset of piscivory, 100–260mm SL subtropical largemouth bass began utilizing more crustaceans than fish. In contrast, temperate M. salmoides consistently fed on fish following the onset of piscivory. Variations in food habits were associated with differences in gape size between temperate and subtropical populations. Temperate largemouth bass had significantly larger gape height (ANCOVA: F = 103.4; df = 1,536; p < 0.001) and width (ANCOVA: F = 47.0; df = 1,536; p < 0.001) than subtropical bass. Although piscivory is a well-known feature of M. salmoides, the ontogeny of piscivory may vary between populations. We hypothesize that interpopulation differences in jaw gape may be related to variations in prey-resource utilization.     

Effects of temperature,salinity and body size on routine metabolism of coastal largemouth bass Micropterus salmoides

Routine metabolism (i.e. standard metabolism plus a low level of activity) of coastal largemouth bass Micropterus salmoides from Mobile‐Tensaw Delta, AL, U.S.A. was examined as a function of temperature (15, 20, 25 and 30° C), salinity (0, 4, 8 and 12) and body mass (range 24–886 g) using flow‐through respirometry. Functionally, a cubic relationship best described the effect of salinity on respiration; the magnitude of these effects increased with temperature and body mass. The best model predicted that specific respiration (mg O₂ g^?1 h^?1) at temperatures >20° C was lowest at salinities of 0·0 and 9·7, and elevated at 3·2 and 12·0; salinity had little to no effect at temperatures ≤20° C. Respiration increased exponentially with temperature, but when compared with previously published respiration rates for M. salmoides from northern latitudes, predicted respiration was higher at cool temperatures and lower at high temperatures. The reduced energetic cost near the isosmotic level (i.e. c. 9) may be an adaptive mechanism to tolerate periods of moderate salinity levels and may help explain why M. salmoides do not flee an area in response to increased salinity. Further, these results suggest that salinity has high energetic costs for coastal populations of M. salmoides and may contribute to the observed slow growth and small maximum size within coastal systems relative to inland freshwater populations.     

Backpack electrofishing effort and imperfect detection: Influence on riverine fish inventories and monitoring

An increased electrofishing sampling effort will increase detection probabilities of riverine fishes. In this study, a repeat‐sampling approach was used in small to medium‐sized Ontario (Canada) rivers to estimate: (i) species‐specific detection probabilities of freshwater fishes, (ii) the number of sampling events required to confidently detect species, and (iii) the power of timed‐search surveys to detect future distribution (or occupancy) declines. Wadeable habitats at 36 sites were sampled with a backpack electrofisher on four separate dates during the summer low‐flow period in 2013 and 2014. Forty‐two species were collected, including three species of conservation concern (American eel Anguilla rostrata Lacépède, 1802, channel darter Percina copelandi Jordan, 1877, northern sunfish Lepomis peltastes Cope, 1870), and two recreationally important species (largemouth bass Micropterus salmoides Lacépède, 1802 and smallmouth bass Micropterus dolomieu Lacépède, 1802). A hierarchical Bayesian modelling approach was used to estimate detection probabilities and site occupancy for 18 species at four levels of effort: 250, 500, 750 and 1,000 s. In all cases, species detection was imperfect. Search effort had a positive effect on estimates of detection probability and site occupancy and the power to detect declines in future distribution. Detection probabilities ranged from 0.11 to 0.66 with an effort of 250 s, and 0.27 to 0.92 with an effort of 1,000 s. For 13 species, detection and power to detect changes in distribution were significantly improved by increasing sampling effort from 250 to 750 s or 1,000 s. For the channel darter and northern sunfish, three replicate sampling visits (of 750 or 1,000 s duration) are recommended for confident detection.