Ecological inferences on invasive carp survival using hydrodynamics and egg drift models.

Bighead carp (Hypophthalmichthys nobilis), silver carp (H. molitrix), black carp (Mylopharyngodon piceus), and grass carp (Ctenopharyngodon idella), are invasive species in North America. However, they hold significant economic importance as food sources in China. The drifting stage of carp eggs has received great attention because egg survival rate is strongly affected by river hydrodynamics. In this study, we explored egg-drift dynamics using computational fluid dynamics (CFD) models to infer potential egg settling zones based on mechanistic criteria from simulated turbulence in the Lower Missouri River. Using an 8-km reach, we simulated flow characteristics with four different discharges, representing 45-3% daily flow exceedance. The CFD results elucidate the highly heterogeneous spatial distribution of flow velocity, flow depth, turbulence kinetic energy (TKE), and the dissipation rate of TKE. The river hydrodynamics were used to determine potential egg settling zones using criteria based on shear velocity, vertical turbulence intensity, and Rouse number. Importantly, we examined the difference between hydrodynamic-inferred settling zones and settling zones predicted using an egg-drift transport model. The results indicate that hydrodynamic inference is useful in determining the 'potential' of egg settling, however, egg drifting paths should be taken into account to improve prediction. Our simulation results also indicate that the river turbulence does not surpass the laboratory-identified threshold to pose a threat to carp eggs.

Movement ecology of diploid and triploid grass carp in a large reservoir and upstream tributaries.

Grass carp Ctenopharyngodon idella, is an herbivorous fish originally brought to North America from Asia in 1963 to control nuisance aquatic vegetation. Since their arrival, detrimental alterations to aquatic ecosystems have sometimes occurred in waterways where they were initially stocked and into which they have escaped. The movements of grass carp from lentic systems into tributaries required for spawning is poorly understood, and understanding environmental conditions associated with upstream migrations may aid in management of the species. We stocked 43 fertile diploid and 43 sterile triploid grass carp implanted with acoustic transmitters into Truman Reservoir, Missouri, USA between January 2017 and October 2018 to characterize movements during spring and summer when spawning conditions occur. Twenty fish (11 diploid/9 triploid) exhibited upstream migration behavior in the Osage River, a major tributary, in 2018 and 2019. Migration primarily occurred in April and May, during high discharge events associated with increasing river stage when water temperatures were between 15 and 28°C. Observed migrations ranged from 3.0-108 river km in length, and six individuals were observed making multiple upstream migrations in one season. Eleven fish initiated upstream migrations while in the lentic main body of the reservoir. These findings provide some evidence for upstream migrations by diploid and triploid grass carp as well both lake and river residents. Evidence of similar upstream migration behavior by both diploid and triploid grass carp suggests that triploids may be suitable surrogates for diploids for study of movement ecology. Removal efforts in tributaries targeting periods of increasing river stage during spring may provide the best opportunity of encountering large concentrations of grass carp.

Developmental ecomorphology of the epibranchial organ of the silver carp, Hypophthalmichthys molitrix.

Silver carp regularly consume and digest particles of food as small as 5 µm. This ability drives their efficient consumption of phytoplankton and because they feed low on the food chain they have an important place in aquaculture worldwide. In North America, where they are considered invasive, silver carp deplete food resources for native species and in so doing occupy increased niche space. Here, we determine the ontogenetic stage and size at which silver carp are morphologically capable of primarily feeding on particles <10 µm. Ecological studies on this species have shown that there is an ontogenetic shift in diet as predominantly zooplanktivorous juveniles later switch to eating much smaller phytoplankton. The occupation of this new trophic niche presents both a metabolic and a mechanical challenge to these fish, since it is unclear how they can efficiently feed on such small particles. We hypothesize that the epibranchial organ (EBO) in silver carp is essential in aggregating these small particles of food, allowing the species to consume mass quantities of tiny particles, thus mitigating metabolic constraints. In this study, we investigate early ontogeny of the EBO in silver carp to determine when this structure achieves the requisite morphology to become functional. We find that at around 80 mm standard length (SL) the EBOs are consistently filled with food, demonstrating that this accumulating organ has become functional. This size corresponds with previous ecological data documenting important shifts in the type of food consumed. While the basic bauplan of the EBO is established very early in ontogeny (by 15 mm SL), multiple waves of histological maturation of muscle, cartilage, gill rakers and epithelium ultimately form the functional structure.

Survival and drifting patterns of grass carp eggs and larvae in response to interactions with flow and sediment in a laboratory flume.

A series of laboratory experiments was conducted to better understand the behavior of grass carp eggs and larvae in moving water in order to develop and implement new strategies for control and prediction of their dispersal and drift at early life stages. Settling velocity and density of a representative sample of eggs were estimated, and three trials of flume experiments with different flow conditions were conducted with live eggs in a temperature-controlled setting with a mobile sediment bed. In these trials, egg and larval stages were continuously analyzed over periods of 80 hours; and eggs and larvae interactions with the flow and sediment bed were monitored and characterized qualitatively and quantitatively. Survival rates were quantified after each trial, highlighting physical causes for increased mortality. Detailed flow analysis was correlated to the observed drifting and swimming behavior of eggs and larvae, to estimate distributions across the water depth, as well as traveling and swimming speeds. Evidence of the influence of mean and turbulent flow in the suspension and transport of eggs are reported, and swimming patterns of larvae at different developmental stages are described. These findings support the development of new strategies for monitoring the spread of grass carp eggs and larvae in rivers, and provide new inputs to predict conditions favorable for spawning and hatching, allowing for mitigation measures at early life stages, which are critical to control their dispersal.

Ontogenetic changes in swimming speed of silver carp, bighead carp, and grass carp larvae: implications for larval dispersal.

Bighead, silver, and grass carps are invasive in the waterways of central North America, and grass carp reproduction in tributaries of the Great Lakes has now been documented. Questions about recruitment potential motivate a need for accurate models of egg and larval dispersal. Quantitative data on swimming behaviors and capabilities during early ontogeny are needed to improve these dispersal models. We measured ontogenetic changes in routine and maximum swimming speeds of bighead, grass, and silver carp larvae. Daily measurements of routine swimming speed were taken for two weeks post-hatch using a still camera and the LARVEL program, a custom image-analysis software. Larval swimming speed was calculated using larval locations in subsequent image frames and time between images. Using an endurance chamber, we determined the maximum swimming speed of larvae (post-gas bladder inflation) for four to eight weeks post-hatch. For all species, larval swimming speeds showed similar trends with respect to ontogeny: increases in maximum speed, and decreases in routine speed. Maximum speeds of bighead and grass carp larvae were similar and generally faster than silver carp larvae. Routine swimming speeds of all larvae were highest before gas bladder inflation, most likely because gas bladder inflation allowed the fish to maintain position without swimming. Downward vertical velocities of pre-gas bladder inflation fish were faster than upward velocities. Among the three species, grass carp larvae had the highest swimming speeds in the pre-gas bladder inflation period, and the lowest speeds in the post-gas bladder inflation period. Knowledge of swimming capability of these species, along with hydraulic characteristics of a river, enables further refinement of models of embryonic and larval drift.

Embryonic and larval development and early behavior in grass carp, Ctenopharyngodon idella: implications for recruitment in rivers.

With recent findings of grass carp Ctenopharyngodon idella in tributaries of the Great Lakes, information on developmental rate and larval behavior is critical to efforts to assess the potential for establishment within the tributaries of that region. In laboratory experiments, grass carp were spawned and eggs and larvae reared at two temperature treatments, one "cold" and one "warm", and tracked for developmental rate, egg size, and behavior. Developmental rate was quantified using Yi's (1988) developmental stages and the cumulative thermal units method. Grass carp had a thermal minimum of 13.5°C for embryonic stages and 13.3°C for larval stages. Egg size was related to temperature and maternal size, with the largest eggs coming from the largest females, and eggs were generally larger in warmer treatments. Young grass carp larvae exhibited upward and downward swimming interspersed with long periods of lying on the bottom. Swimming capacity increased with ontogeny, and larvae were capable of horizontal swimming and position holding with gas bladder emergence. Developmental rates, behavior, and egg attributes can be used in combination with physical parameters of a river to assess the risk that grass carp are capable of reproduction and recruitment in rivers.

Aspects of embryonic and larval development in bighead carp Hypophthalmichthys nobilis and silver carp Hypophthalmichthys molitrix.

As bighead carp Hypophthalmichthysnobilis 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.

Microsatellite genetic diversity and differentiation of native and introduced grass carp populations in three continents.

Grass carp (Ctenopharyngodon idella), a freshwater species native to China, has been introduced to about 100 countries/regions and poses both biological and environmental challenges to the receiving ecosystems. In this study, we analyzed genetic variation in grass carp from three introduced river systems (Mississippi River Basin in US, Danube River in Hungary, and Tone River in Japan) as well as its native ranges (Yangtze, Pearl, and Amur Rivers) in China using 21 novel microsatellite loci. The allelic richness, observed heterozygosity, and within-population gene diversity were found to be lower in the introduced populations than in the native populations, presumably due to the small founder population size of the former. Significant genetic differentiation was found between all pairwise populations from different rivers. Both principal component analysis and Bayesian clustering analysis revealed obvious genetic distinction between the native and introduced populations. Interestingly, genetic bottlenecks were detected in the Hungarian and Japanese grass carp populations, but not in the North American population, suggesting that the Mississippi River Basin grass carp has experienced rapid population expansion with potential genetic diversification during the half-century since its introduction. Consequently, the combined forces of the founder effect, introduction history, and rapid population expansion help explaining the observed patterns of genetic diversity within and among both native and introduced populations of the grass carp.