Trophic reorganization of native planktivorous fishes at different density extremes of bigheaded carps in the Illinois and Mississippi rivers,USA

The widespread introduction of non-native fishes has contributed to freshwater ecosystems being considered among the most altered ecosystems globally. Of particular concern are invasive planktivorous fishes (e.g., silver carp Hypophthalmichthys molitrix and bighead carp H. nobilis, collectively known as bigheaded carps) that have the potential to modify basal food web structure and compete for planktonic resources with native planktivores and young-of-year fishes. Bigheaded carps have proliferated throughout the Mississippi River basin, creating an outsized potential for resource competition with native fishes. Studies have showed niche overlap between bigheaded carps and native planktivores is generally high but that overlap varies among rivers. Importantly, niche overlap has not been assessed for density extremes of bigheaded carps within a river to determine whether trophic niches changed as a result of the invasion. The objectives of this study were to determine whether (1) silver and/or bighead carps share a similar isotopic niche with four native planktivores, and (2) that association varies ecologically (i.e., low- and high-densities of bigheaded carps) and spatially (i.e., between rivers). Our results generally show high trophic overlap among species, suggesting potential direct resource competition. Niche overlap was higher in study reaches with low densities of bigheaded carps compared to reaches with high densities, presumably due to intense resource competition and limiting of resources under high densities of bigheaded carps. Across density extremes, trophic reorganization by bigmouth buffalo (Ictiobus cyprinellus) was divergent from other native planktivores. Species-specific responses may be due to subtle differences in feeding strategies, degree of planktivory, food selectivity, and correlated food size distributions.

     

Reproductive condition and occurrence of intersex in bighead carp and silver carp in the Missouri River

Little is known about the reproductive biology of the exotic bighead carp Hypophthalmichthys nobilis and silver carp Hypophthalmichthys molitrix in the Missouri River. In order to fill this gap in understanding, herein is described the reproductive condition of these Asian carps. Evidence is presented which indicates that bighead and silver carp in the Missouri River have a protracted spawning period that extends from early spring through fall and some individual bighead and silver carp are spawning multiple times during a reproductive season. Although bighead and silver carps are successfully maturing and spawning in the Missouri River some reproductive abnormalities such as intersex, atresia, and sterility were observed. Knowledge of the reproductive activity of these invasive carps may be useful to resource managers tasked with their control. Furthermore, the reproductive abnormalities observed should be considered when evaluating the environmental condition of the Missouri River relative to supporting a healthy fish fauna.     

Impacts of two biomanipulation fishes stocked in a large pen on the plankton abundance and water quality during a period of phytoplankton seasonal succession

Silver and bighead carp were stocked in a large pen to control the nuisance cyanobacterial blooms in Meiliang Bay of Lake Taihu. Plankton abundance and water quality were investigated about once a week from 9 May to 7 July in 2005. Biomass of both total crustacean zooplankton and cladocerans was significantly suppressed by the predation of pen-cultured fishes. There was a significant negative correlation between the N:P weight ratio and phytoplankton biomass. The size-selective predation by the two carps had no effect on the biomass of green alga Ulothrix sp. It may be attributed to the low fish stocking density (less than 40 g m⁻³) before June. When Microcystis dominated in the water of fish pen, the pen-cultured carps effectively suppressed the biomass of Microcystis, as indicated by the significant decline of chlorophyll a in the >38 μm fractions of the fish pen. Based on the results of our experiment and previous other studies, we conclude that silver and bighead carp are two efficient biomanipulation tools to control cyanobacterial (Microcystis) blooms in the tropical/subtropical eutrophic waters. Moreover, we should maintain an enough stocking density for an effective control of phytoplankton biomass.     

Swimming performance of adult bighead carp Hypophthalmichthys nobilis (Richardson, 1845) and silver carp H. molitrix (Valenciennes, 1844)

Although the movement of invasive bighead carp (Hypophthalmichthys nobilis) and silver carp (H. molitrix) in the Upper Mississippi River system is dependent on their ability to swim through its numerous lock‐and‐dams, the swimming performance of adults of these species is at present unknown. Using a large (2,935‐L) mobile swim tunnel, the swimming performance of adult bighead and adult silver carp was quantified at water velocities that challenged them to exhibit either prolonged and/or burst swimming (76–244 cm/s) with fatigue times of less than 10 min. Simple log‐linear models best described the relative swim speed to fatigue relationships for both species. Under these conditions, the swimming performances of adult bighead and silver carp were similar to several species of adult fishes native to the Mississippi River system, but relatively low (<3 total body lengths per second, TL/s) compared to previously studied juveniles and sub‐adult bigheaded carps (3–15 TL/s). The decline in endurance with water velocity was three times greater in bighead carp (slope = ?2.98) than in silver carp (slope = ?1.01) and the predictive ability of the bighead model was appreciably better than the silver carp model. The differences in adult swimming performance between the two species were coincident with behavioral differences (e.g. breaching in silver carp but not in bighead carp). The swimming performance data of adult bighead and silver carp can now be used to evaluate whether their passage through manmade river structures including the gates of lock‐and‐dams in the Upper Mississippi River might be reduced.     

Bigheaded carps (<Emphasis Type="Italic">Hypophthalmichthys</Emphasis> spp.) at the edge of their invaded range: using hydroacoustics to assess population parameters and the efficacy of harvest as a control strategy in a large North American river

The threat posed by bigheaded carps (Hypophthalmichthys spp.) to novel ecosystems has focused efforts on preventing further range expansion; upstream progression in the Illinois River is a major concern due to its connection with the uninvaded Great Lakes. In addition to an electric barrier system, commercial harvest of silver carp (H. molitrix) and bighead carp (H. nobilis) in the upper river is intended to reduce propagule pressure and prevent range expansion. To quantify demographics and evaluate harvest efficacy, the upper river was sampled between 2012 and 2015 using mobile hydroacoustic methods. Reach-specific densities, size structures and species compositions varied interannually but the advancing population was characterized longitudinally as small-bodied, silver carp-dominated at the highest densities downstream, shifting to large-bodied, bighead carp-dominated at the low-density population front. The use of hydroacoustic sampling for harvest evaluation was validated in backwater lakes; there was a significant positive correlation between density estimates and the corresponding harvest catch-per-unit-effort of bigheaded carps. Localized densities of bigheaded carps were reduced by up to 64.4 % immediately post-harvest but generally rebounded within weeks. However, annual sampling of the entire upper river indicated that density of bigheaded carps decreased by over 40 % (between 2012 and 2013) and subsequently remained stable (between 2013 and 2014). The annual harvest of bigheaded carps increased during this period (from 45,192 to 102,453 individuals), in years of contrasting discharge conditions. At this spatiotemporal scale, harvest appears to have contributed to initial reduction, and subsequent maintenance of, bigheaded carps density levels, but discharge likely plays an important role (e.g., through immigration) in determining the extent of its impact. Mobile hydroacoustic sampling enabled robust quantification of the population over varying spatial scales and density gradients, highlighting the potential of this approach as an assessment tool for invasive fishes in riverine environments.     

Seasonal variations in stable isotope ratios of two biomanipulation fishes and seston in a large pen culture in hypereutrophic Meiliang Bay, Lake Taihu

This paper reports on seasonal changes in stable carbon and nitrogen isotope ratios of seston and muscle tissue of silver carp and bighead carp during 2004 and 2005, focusing primarily on the carbon sources and trophic relationships among phytoplankton, zooplankton and silver carp and bighead carp in a large fish pen of Meiliang Bay (Lake Taihu, China). δ¹³C showed a minimal value in March 2005 and a maximal value in August 2005 in seston both inside and outside the pen, whereas δ¹⁵N of seston showed the minimum in winter and the maximum during algal blooms. A positive correlation between δ¹³C of silver carp and that of seston suggested that temporal variation of δ¹³C in seston was preserved in fish via the food chain. The differences of δ¹³C among seston, zooplankton and muscle tissue of silver carp and bighead carp ranged only 0.2–1.7%, indicating that plankton production was the primary food source of filter-feeding fishes. According to a mass balance model, we estimated that the contributions of zooplankton to the diets of silver carp and bighead carp were 45.7% and 54.3%, respectively, based on the δ¹⁵N values of zooplankton and planktivorous fishes.     

Performance of different fish species for controlling weeds in rainfed lowland rice field

A field experiment was conducted during wet season to evaluate the performance of different fish species for biocontrol of weeds in rainfed waterlogged rice fields with fingerlings of three exotic carps – grass carp, Ctenopharyngodon idella (Val.) (Cyprinidae, Cypriniformes), silver barb, Puntius gonionotus (Bleeker) (Cyprinidae, Cypriniformes) and common carp, Cyprinus carpio (L.) (Cyprinidae, Cypriniformes) – and three Indian major carps – rohu, Labeo rohita (Ham.) (Cyprinidae, Cypriniformes), catla, Catla catla (Ham.) (Cyprinidae, Cypriniformes), mrigal, Cirrhinus mrigala (Ham.) (Cyprinidae, Cypriniformes). A total of 13 major weeds under the categories of grassy, sedges, broadleaf and aquatic weeds were observed in the rice fields. Grass carp reduced maximum weed biomass (weed control efficiency [WCE] 63% at 60 days after transplanting [DAT] and 62% at 100 DAT) followed by silver barb and common carp. Among the Indian carps, only rohu was effective in control of weeds (WCE, 23% at 60 DAT). The grain yield of rice (variety Varshadhan) slightly increased (4.2–4.5 t/ha), but straw yield was significantly higher (10.2–10.7 t/ha) under rice-fish farming. Fish yield was significantly higher in exotic carps (270–288 kg/ha/90days) due to higher specific growth rates (1.8–2% body weight/day). The study indicated that exotic carps (grass carp, silver barb and common carp in order) were more effective than Indian carps for control of weed in rainfed lowland rice fields and among the Indian carps, rohu showed potential for weed control.     

Feeding at different plankton densities alters invasive bighead carp (<Emphasis Type="Italic">Hypophthalmichthys nobilis</Emphasis>) growth and zooplankton species composition

Invasive Asian carps Hypophthalmichthys spp. are an ecological threat to non-native aquatic ecosystems throughout the world, and are poised to enter the Laurentian Great Lakes. Little is known about how these filter-feeding planktivores grow and impact zooplankton communities in mesotrophic to oligotrophic systems like the Great Lakes. Our purpose was to determine how different plankton densities affect bighead carp H. nobilis biomass and how bighead carp affect zooplankton species composition. We conducted a 37-day indoor mesocosm experiment (volume = 678 l) with high and low plankton treatments (zooplankton dry mass ≈ 1,900 and 700 μg l⁻¹; chlorophyll a = 25 and 14 μg l⁻¹, respectively) in the presence and absence of juvenile bighead carp (mean = 5.0 g, 8.5 cm). Carp lost weight in the low plankton treatment and gained weight in the high plankton treatment, suggesting that food availability may be a limiting factor to bighead carp growth in regions of low plankton densities. In the presence of carp, zooplankton shifted from Daphnia to copepod dominance, while in the absence of carp, Daphnia remained dominant. Chydorids and ostracods increased in the presence of carp, but only in the low plankton treatment, suggesting that the impact of bighead carp on zooplankton species composition may vary with zooplankton density. Chlorophyll was higher in the absence of carp than in the presence. Chlorophyll and zooplankton densities in many Great Lakes ecosystems are substantially lower than our low treatment conditions, and thus our results suggest that Asian carp establishment in these regions may be unlikely. Handling editor: S. Declerck     

Tag retention and survival of juvenile bighead carp implanted with a dummy acoustic tag at three temperatures

Bighead carp Hypophthalmichthys nobilis and silver carp Hypophthalmichthys molitrix (together, the bigheaded carps) are invasive fishes in North America that have resulted in substantial negative effects on native fish communities and aquatic ecosystems. Movement and behavior of adult bigheaded carps has been studied previously using telemetry, while similar studies with juvenile bigheaded carps have yet to be attempted. Recent technological advances in telemetry transmitters has increased the availability of tags sufficiently small enough to implant in juvenile carps. However, the effects of surgical implantation of telemetry tags on juvenile bigheaded carps have not been evaluated. We determined tag retention and survival associated with surgical implantation of acoustic telemetry tags into juvenile bighead carp (range 128–152 mm total length) at three temperatures (13, 18, and 23°C). In addition, we assessed the effect of surgically implanted transmitters on the fitness, defined as changes in weight or critical swimming speed, of carp implanted with transmitters. Survival was high among tagged fish (85%) with 47% of tags retained at the conclusion of the 45‐day study. No substantial decline in fitness of the fish was observed in tagged fish compared to untagged fish.     

Comparison of the trophic niches between two planktivorous fishes in two large lakes using stable isotope analysis

This paper studied the sources of food and the trophic levels of silver and bighead carps using stable isotopes (δ¹³C and δ¹⁵N) analysis in Lake Qiandao and Lake Taihu. The δ¹³C values of POM and phytoplankton indicated that Particulate Organic Matter (POM) is of terrestrial origin in LQ and phytoplankton origin in LT. The different signature of δ¹⁵N of phytoplankton and POM also showed the different extent of anthropogenic impacts on the two lakes, with higher impacts in LT than in LQ. In LQ the trophic levels of silver and bighead carps were significantly different (2.48 ± 0.19 and 2.66 ± 0.19, respectively), while in LT the difference was not significant (2.15 ± 1.54 and 2.36 ± 0.38, respectively). Using a mixing model, we found that silver and bighead carps consumed a major proportion on phytoplankton and/or POM in both lakes with higher contribution of phytoplankton in more eutrophic lake. Silver and bighead carps had clear differentiation in food resources with silver carp more on POM and bighead more on zooplankton in deep and mesotrophic lakes. Contrary to this, both carps fed primarily on phytoplankton (and POM) and may have more niche overlaps in shallow and eutrophic lakes.     

Preliminary evidence of delayed spawning and suppressed larval growth and condition of the major carps in the Yangtze River below the Three Gorges Dam

Commercial catches of the ‘major carps’ (grass carp Ctenopharyngodon idella, silver carp Hypophthalmichthys molitrix, black carp Mylopharyngodon piceus and bighead carp Aristichthys nobilis) in the middle and lower reaches of the Yangtze River have declined precipitously since construction of the Three Gorges Dam (TGD). To assess specific environmental effects, young-of-the-year major carps were collected below TGD in the Yangtze River at Jianli (larvae) and E’zhou (juveniles) and in the Dongting Lake (juveniles) during June–August 2007. Lapillar otoliths were analyzed to determine early growth. There were no discernible growth increments in otoliths for a majority of the larvae (38 of 63 grass carp and 40 of 47 silver carp), while increments were evident in otoliths formed in the larval stage for all the juveniles, indicating that larvae without discernible increments failed to recruit to the juvenile stage. Back-calculated hatch dates of major carps were between 31 May and 24 July. The initiation of the spawning season was delayed about 1 month compared to pre-TGD records. Larval growth rates, as reflected by otolith-increment width, were lower at Dongting Lake (closer to TGD) than at E’zhou (farther from TGD), indicating that TGD-moderated effects on early growth and development of carps are more apparent for fishes hatched nearer to the dam. The delay in spawning and decreased early growth may partly explain the recent decline of the major carp resources in the middle and lower Yangtze River.     

In situ study on effect of food competition on diet shifts and growth of silver and bighead carps in large biomanipulation fish pens in Meiliang Bay, Lake Taihu

Silver and bighead carps were cultured in large fish pens to reduce the risks of cyanobacterial bloom outbreaks in Meiliang Bay, Lake Tauhu in 2004 and 2005. Diet compositions and growth rates of the carps were studied from April to November each year. Both carp species fed mainly on zooplankton (>50% in diet) in 2004 when competition was low, but selected more phytoplankton in 2005 when competition was high. Silver carp had a broader diet breadth than did bighead carp. Higher densities and fewer food resources increased diet breadths but decreased the diet overlap in both types of carps. It can be predicted that silver and bighead carps would be released from diet competition and shift to feed mainly on zooplankton at low densities, decreasing the efficiency of controlling cyanobacterial blooms. Conclusively, when silver and bighead carps are used to control cyanobacterial blooms, a sufficiently high stocking density is very important for a successful practice.     

Relationship between meteorological conditions and natural reproduction behavior of the four famous Chinese carps

Known as the four famous carps in China, black carp (Mylopharyngodon piceus), grass carp (Ctenopharyngodon idella), silver carp (Hypophthalmichthys molitrix) and bighead carp (Hypophthalmichthys nobilis) are commercially important fish species with a high production in China. To reveal the relationships between meteorological conditions and the natural spawning behavior of these fishes, we considered six meteorological factors (including wind, rainfall, temperature, air pressure, sunshine hours and humidity) and the weather conditions in 80 spawning events. The results showed that the spawning activities were more likely to be activated in consecutive rainy days or days when weather changed drastically, while the spawning activities showed no tendency for the weather type except consecutive rainy days. Our analyses also showed that the average rainfall in the initial spawning days is higher than that in the spawning time windows (the time from the earliest spawning activity to the latest spawning activity), while other meteorological elements remained at more or less the same values in both time periods; spawning activity tended to happen in days when the average air pressure was going down or the average temperature going up, while the other meteorological elements showed no consistent tendency.     

Temperature dependent effects of carbon dioxide on avoidance behaviors in bigheaded carps

Effective behavioral deterrents are needed to prevent aquatic invasive species from entering novel ecosystems. One deterrent strategy that shows promise is elevated carbon dioxide (CO₂) concentrations in water which can alter the behavior of freshwater fishes, including invasive bigheaded carps (Hypophthalmichthys spp.). However, few studies have evaluated behavioral responses to elevated CO₂ concentrations at different water temperatures. The objective of this study was to quantify CO₂ concentrations needed to achieve avoidance (voluntary response) and narcosis (involuntary response observed by loss of equilibrium) behaviors in silver carp (H. molitrix) and bighead carp (H. nobilis) at 5, 15, and 25 °C. Overall, silver carp and bighead carp displayed avoidance and narcosis behaviors to CO₂ at each water temperature, however bighead carp responded at higher CO₂ concentrations than silver carp. Behavioral avoidance and narcosis were observed at approximately 40% lower CO₂ concentrations in 5 °C water relative to 25 °C suggesting considerable influence of water temperature on a CO₂ stimulus for both species. Results indicate that fluctuating water temperature (e.g., spatial and temporal variation across management sites) can influence how fish respond to elevated CO₂, and may usefully be considered when applying CO₂ as a behavioral deterrent.     

Persistence of DNA in Carcasses,Slime and Avian Feces May Affect Interpretation of Environmental DNA Data

The prevention of non-indigenous aquatic invasive species spreading into new areas is a goal of many resource managers. New techniques have been developed to survey for species that are difficult to capture with conventional gears that involve the detection of their DNA in water samples (eDNA). This technique is currently used to track the invasion of bigheaded carps (silver carp and bighead carp; Hypophthalmichthys molitrix and H. nobilis) in the Chicago Area Waterway System and Upper Mississippi River. In both systems DNA has been detected from silver carp without the capture of a live fish, which has led to some uncertainty about the source of the DNA. The potential contribution to eDNA by vectors and fomites has not been explored. Because barges move from areas with a high abundance of bigheaded carps to areas monitored for the potential presence of silver carp, we used juvenile silver carp to simulate the barge transport of dead bigheaded carp carcasses, slime residue, and predator feces to determine the potential of these sources to supply DNA to uninhabited waters where it could be detected and misinterpreted as indicative of the presence of live bigheaded carp. Our results indicate that all three vectors are feasible sources of detectable eDNA for at least one month after their deposition. This suggests that current monitoring programs must consider alternative vectors of DNA in the environment and consider alternative strategies to minimize the detection of DNA not directly released from live bigheaded carps.     

Modelling bioenergetic and population-level impacts of invasive bigheaded carps (Hypophthalmichthys spp.) on native paddlefish (Polyodon spathula) in backwaters of the lower Mississippi River

1. While invasions of large rivers by exotic fish species are well documented, assessing actual or potential impacts on native species is a challenge. Rapid assessments may be possible through the application of a combination of bioenergetic and population dynamic models.
2. Paddlefish (Polyodon spathula) is a native species in the central USA with a history of population decline due to waterway development and overharvesting for roe. It is not known whether paddlefish are impacted by resource competition from invasive bigheaded carp populations, including silver (Hypophthalmichthys molitrix) and bighead carp (Hypophthalmichthys nobilis), which have expanded dramatically in the Mississippi River.
3. We used bioenergetic models to project the potential impact of invasive silver and bighead carp on zooplankton density and paddlefish somatic growth in backwater habitat. Bioenergetic outputs were translated to impacts on fecundity, becoming inputs for 50-year metapopulation simulations of backwater habitat connected to the main-stem Mississippi River by episodic flood events.
4. Competition with carp reduced growth and increased the risk of population decline for paddlefish. Impacts increased disproportionately with increased carp abundance and were further exacerbated in scenarios with increased diet overlap or decreased zooplankton abundance.
5. We also analysed paddlefish condition data collected at sites near the lower Mississippi River with varying histories of carp invasion. These data give credence to the bioenergetic model output; paddlefish had reduced body condition at sites with long-established, high-density carp populations.
6. We conclude that invasive bigheaded carps have great potential to reduce paddlefish growth, fecundity, and abundance. The pairing of bioenergetics and population models is likely to be broadly useful in assessing the risks posed by other invasive species.

     

Long-term changes in fish community structure in relation to the establishment of Asian carps in a large floodplain river

The Upper Mississippi River System, including the Illinois River, has been invaded by a number of nonnative species including silver carp Hypophthalmichthys molitirx and bighead carp H. nobilis, collectively referred to here as Asian carps. Silver carp densities in the Illinois River have increased dramatically and now represent some of the highest densities of wild silver carp anywhere in the world. Asian carps have the potential to alter existing ecosystems by consuming planktonic resources and therefore, could have the ability to alter existing fish communities as most fishes are dependent on planktonic resources during early development. However, identifying the relationship of fish community structure to the establishment of Asian carps has yet to be thoroughly investigated. Using long-term fish community data collected by the Upper Mississippi River Restoration’s Long Term Resource Monitoring element, we investigate changes in fish community structure pre- and post-establishment of Asian carps. Significant differences in the pre- and post-establishment communities were observed for the majority of gears and habitats. Species contributing to changes between establishment periods included most sportfish species and catostomids, which were less abundant post-establishment of Asian carps, while shortnose gar, grass carp, and emerald shiner were more abundant. While our analyses show Asian carps are likely contributing to major differences in fish community structure, future research and long-term monitoring should investigate the mechanisms and interactions responsible for community changes as well as identifying any potential concurrent or confounding factors such as changes in river hydrology or sedimentation.     

Aspects of Embryonic and Larval Development in Bighead Carp Hypophthalmichthys nobilis and Silver Carp Hypophthalmichthys molitrix

As bighead carp Hypophthalmichthys nobilis and silver carp H . molitrix (the bigheaded carps) are poised to enter the Laurentian Great Lakes and potentially damage the region’s economically important fishery, information on developmental rates and behaviors of carps is critical to assessing their ability to establish sustainable populations within the Great Lakes basin. In laboratory experiments, the embryonic and larval developmental rates, size, and behaviors of bigheaded carp were tracked at two temperature treatments, one “cold” and one “warm”. Developmental rates were computed using previously described stages of development and the cumulative thermal unit method. Both species have similar thermal requirements, with a minimum developmental temperature for embryonic stages of 12.1° C for silver carp and 12.9° C for bighead carp, and 13.3° C for silver carp larval stages and 13.4° C for bighead carp larval stages. Egg size differed among species and temperature treatments, as egg size was larger in bighead carp, and “warm" temperature treatments. The larvae started robust upwards vertical swimming immediately after hatching, interspersed with intervals of sinking. Vertical swimming tubes were used to measure water column distribution, and ascent and descent rates of vertically swimming fish. Water column distribution and ascent and descent rates changed with ontogeny. Water column distribution also showed some diel periodicity. Developmental rates, size, and behaviors contribute to the drift distance needed to fulfill the early life history requirements of bigheaded carps and can be used in conjunction with transport information to assess invasibility of a river.