Development of microsatellite markers for muskellunge (Esox masquinongy) and cross‐species amplification in two other esocids

We report the development of seven polymorphic microsatellite loci in muskellunge (Esox masquinongy) using an unenriched subgenomic library. Polymorphic loci exhibited 2–11 alleles with observed heterozygosities ranging from 0.190 to 0.917 (n = 24). All seven loci amplified by their respective primer pairs resulted in monomorphic products in northern pike (E. lucius) whereas three loci amplified monomorphic products in grass pickerel (E. americanus americanus); these results imply conservation of flanking sequence but loss of polymorphism between these closely related species. Only one of six microsatellite primers developed in a previous study in northern pike amplified polymorphic products in muskellunge.     

History of muskellunge management in Nebraska

Muskellunge, Esox masquinongy, were first introduced to Nebraska waters in 1958. By 1967 anglers had caught the first muskellunge from Nebraska waters. Hybrid muskellunge, Esox masquinongy × E. lucius, culture and stocking started in 1976. Harvest regulations for muskellunge and hybrid muskellunge were first established in 1977 with a daily bag limit of one muskellunge or hybrid muskellunge of at least 762 mm (30 inches) in length. In 2000 the minimum length limit for muskellunge and hybrid muskellunge was increased to 1016 mm (40 inches). Hybrid muskellunge have been phased out, none have been stocked in Nebraska waters since 2003. Since 1998 muskellunge fingerlings have been held in Nebraska hatcheries until age-1 before stocking. Yearling muskellunge are raised on minnows and other small fish in Nebraska hatcheries and stocking success apparently is very good. Management and research of Nebraska’s muskellunge and hybrid muskellunge fisheries has been limited to routine sampling that has documented the presence of those fish in waters where they were stocked. Fishable populations of muskellunge can now be found in nine Nebraska waters across the state, but most Nebraska anglers are relatively uneducated in fishing techniques for muskellunge, identification, and proper handling.     

An examination of Minnesota’s muskellunge waters

Synopsis We examined Minnesota’s muskellunge, Esox masquinongy, populations using various data sets including spring trap net assessments and angler diary surveys. We confronted a potentially problematic data set that had a large number of zero catches by fitting logistic models to the presence/absence data set and linear models to the subset of data that included catches with one fish or more. Currently, 105 lakes have been identified as muskellunge waters, of which 61 lakes have been created and maintained by stocking. Although the proportion of successful anglers has increased over time, catch rates have remained the same. It appears from analysis of trap net catches that the abundance of 102 cm (40 in) and larger muskellunge has been increasing over time. Both trap net and angler data provide some indications that size of muskellunge caught has also increased over time. Angler-harvested muskellunge averaged 11 years of age and 115 cm (45 in) total length. For Minnesota waters, muskellunge ultimate length averaged 134 cm (53 in) for females and 119 cm (47 in) for males. All evidence indicates a successful management program.     

Expulsion of miniature radio transmitters along with eggs of muskellunge and northern pike––a new method for locating critical spawning habitat

Identification and protection of critical spawning habitat for muskellunge Esox masquinongy and northern pike Esox lucius is important for preserving the reproductive potential of both species. In this study, we implanted miniature radio transmitters through the oviduct into the egg masses of female muskellunge and northern pike just prior to spawning. This non-surgical procedure was a novel approach for identifying spawning sites when transmitters were expelled with the eggs during egg deposition. Preliminary studies in three lakes showed that muskellunge and northern pike deposited many of the transmitters in likely spawning habitat. An inability to find eggs limited our validation of this method, but nevertheless, a relatively high proportion (70%) of northern pike larger than 690 mm (27.2 inches) expelled transmitters in a previously known spawning area in Willow Lake, Minnesota. Shoreline vegetation in that area consisted primarily of sedges Carex spp., and the adjacent water was shallow with substrate consisting of large mats of water bulrush Scirpus subterminalis. A lower proportion (50%) of muskellunge expelled transmitters in Elk Lake, Minnesota. Water depth at likely spawning sites averaged 1.1 m (3.6 feet) and vegetative cover was variable, but Chara spp. was common to most sites. In Moose Lake, Minnesota, containing sympatric populations of muskellunge and northern pike, 60% of muskellunge and 90% of pike expelled transmitters. Chara spp. beds were the predominant substrate where transmitters were expelled in Moose Lake, but the two species deposited transmitters on deepwater bars (3.7–5.2 m) in addition to shallow near-shore habitat. These results suggest more flexibility in depths used for spawning than typically reported for muskellunge and northern pike.     

Effects of angling with a single-hook and live bait on muskellunge survival

Use of live bait for angling of muskellunge, Esox masquinongy, is popular in Wisconsin. A traditional method utilizes a large hook through the bait fish’s snout, which requires the muskellunge to swallow the bait prior to hook set. Adult muskellunge (>76 cm; 30 in) were held in lined hatchery ponds and caught while fishing with live bait on 10/0 size single hooks. The leader was cut and the muskellunge was released when hooked in the stomach. Survival was monitored for up to 1 year. No immediate ( <24 h) mortality occurred. However, 22% of hooked muskellunge died within 50 days and 83% died within 1 year. Necropsies revealed extensive trauma to the stomach and other organs from hooks, along with systemic bacterial infections. Highest mortality on both hooked and control fish occurred over winter through spring. This peak mortality may be associated with natural stressors that occur during the spring spawning period. Mortality rates observed in this study are considered unacceptable for trophy management of muskellunge. Although use of live bait for muskellunge is traditional in Wisconsin, terminal tackle such as quick-strike rigs that hook fish in the mouth or buccal cavity should enhance the chances a released muskellunge will survive.     

Comparison of mouth morphology and prey size selection among three esocid taxa

Aquatic organisms, especially fishes, exhibit exceptional diversity in mouth morphology and this variation has been shown to influence foraging patterns. We compared mouth morphology among muskellunge Esox masquinongy, northern pike Esox lucius and their hybrid, tiger muskellunge E. masquinongy x E. lucius. Head and mouth size among the three taxa were similar for juveniles (<400 mm total length), but diverged with increasing length, being greater for northern pike than muskellunge. Tiger muskellunge had a head and mouth size intermediate to the two, but more similar to northern pike than muskellunge. Morphological differences among taxa were related to data examining prey size selection in laboratory and field experiments. In the laboratory, northern pike selected prey that were smaller than their maximum mouth width (widest point between outside corners of mouth), tiger muskellunge selected larger prey, and muskellunge size-selection was intermediate between the other two taxa. Among the three esocids, muskellunge had the smallest increase in handling time with increasing prey body depth relative to predator mouth width. In a common garden field experiment in three lakes containing mainly deep-bodied prey, results generally followed morphological patterns, with northern pike selecting larger prey compared to muskellunge. Although morphology predicted most of the variation in greatest body depth of prey consumed, the best predictor of prey size was a model that included predator mouth width, taxon, and interaction. Information comparing prey size selection among esocid taxa is useful for understanding how to manage esocid populations based on system-specific prey characteristics and also for understanding how variations in morphological characteristics of apex predators can influence prey vulnerability and ecosystem structure.     

Influence of cisco (Coregonus artedi,Lesueur) on muskellunge (Esox masquinongy,Mitchill) mean length,population size structure,and maximum size in northern Wisconsin lakes

Population size structure and maximum size of managed sportfish populations are dictated by abiotic, biotic, ecosystem, and anthropogenic influences. In their native ranges of northern Wisconsin, muskellunge (Esox masquinongy) and cisco (Coregonus artedi) are co-adapted cool- and cold-water species where cisco presence may influence population size structure and maximum size of muskellunge. We tested whether muskellunge size structure indices (length-frequency distributions, proportional size distribution), mean length, and mean maximum length of muskellunge differed when cisco were present or absent in Ceded Territory of Wisconsin (CTWI) lakes during 2015–2018. Cisco presence had a positive influence on size structure and mean length of individual muskellunge within populations. In contrast, cisco presence had no influence on the mean maximum length of muskellunge observed in CTWI populations suggesting that other factors may be better predictors of this metric than cisco presence. In cisco lakes, mean muskellunge length was negatively correlated with mean cisco length suggesting that gape limitation may be a factor influencing population size structure and individual growth rates. Therefore, cisco populations with primarily large individuals may be unavailable to muskellunge as forage. Our results suggest that cisco are an important forage species for some aspects of muskellunge population ecology; however, other factors may also contribute to muskellunge population size structure and maximum size outcomes. As such, conservation of remaining cisco populations in Wisconsin is critical because they influence muskellunge population ecology in lakes where the species coexist. Future research is needed to better understand the interactions of cisco, abiotic and biotic factors, and anthropogenic influences on muskellunge growth dynamics.     

Characteristics of Ontario muskellunge (Esox masquinongy) fisheries based on volunteer angler diary information

This paper consolidates and summarizes information on Ontario muskellunge, Esox masquinongy, sport fisheries derived from angler diary programs sponsored by the Ontario Ministry of Natural Resources (OMNR) and Muskies Canada Incorporated (MCI) from 1979 to 2004. Interest in muskellunge as a sport fish has increased substantially over the past 10–15 years. Muskellunge catches were found to be highly correlated with angling effort. Catch rates, expressed in terms of catch-per-unit-of-effort (CUE), have improved over the past decade to the point where, in 2001, the provincial CUE was 0.119. The long term catch rate is 0.069. This improvement in angling quality is attributed to new minimum size limit regulations and increased catch-and-release angling practices. Over the period from 1979–2004, release rates by muskellunge anglers have averaged 94%. Based on a sample size of 9,499 fish, the mean size of angled muskellunge over the past 26 years was 37.0 inches (94 cm). Numerous fish exceeding 50 inches (127 cm) are angled from Ontario waters each year. The incidence of lymphosarcoma, a highly contagious, malignant blood cancer, has averaged only 2% since 1979. Based on an analysis of this information, Ontario’s muskellunge fisheries appear to be stable and sustainable. Volunteer angler diary programs provide an accurate and cost-effective means to monitor the status of muskellunge fisheries in Ontario.     

Induction of gynogenesis in muskellunge with irradiated sperm of yellow perch proves diploid muskellunge male homogamety

Diploid gynogenesis was induced in muskellunge Esox masquinongy using UV-irradiated muskellunge sperm as the first step in producing monosex females. In this approach, we have to rely on negative controls as an indirect reference for sperm genetic material destruction. In the first experiment, equal proportions of gynogenetic females and males were produced. Negative controls, UV-irradiated sperm without heat shock, yielded some normal hatching larvae, described as spontaneous diploids. In the second experiment, muskellunge eggs were activated using sperm from yellow perch. Because hybrids between these species are not viable, we produced unambiguous gynogens. When UV-irradiated yellow perch sperm was used to inseminate muskellunge eggs, haploids resulted (22.5% +/- 2.8% survival to the eyed stage). To produce diploid gynogens, a heat shock of 31 degrees C was applied to inseminated eggs 20 min after activation for a duration of 6 min. This process yielded several hundreds of gynogens for rearing. Several treatments of masculinizing hormone, 17alpha-methyltestosterone (MT), were carried out. Fish were dissected and gonads examined histologically for sex determination. Gynogens produced using yellow-perch sperm confirmed the presence of males in the control group, whereas the MT bath treatment (400 microg/liter) resulted in the production of fish with ovotestis. These results provide evidence for male homogamety in muskellunge and imply that a change of strategy is needed to produce monosex populations.     

Development, implementation, and evaluation of an international muskellunge management strategy for the upper St Lawrence River

The muskellunge, Esox masquinongy, fishery in the St Lawrence River is believed to have declined significantly from historical levels and reached critically low levels during the 1970s. Over-exploitation caused by liberal angling regulations, and loss and alteration of critical spawning and nursery habitat probably contributed to this decline. In 1980, a St Lawrence River Muskellunge Management Work Group comprising resource managers and several advisors, including E.J. Crossman, to whom this symposium is dedicated, was created to address research and management needs. A trophy muskellunge management strategy was implemented including more restrictive harvest regulations, public education promoting “catch and release”, and protection of spawning and nursery habitats. Age and growth information obtained from cleithra analysis indicated the need for increased size limits to adequately protect spawning stocks. Research efforts have developed a biological information base and monitoring tools to guide management decisions and evaluate responses. Over 100 spawning and nursery locations have been identified in US and Canadian waters leading to improved protection of critical habitats. An angler diary program shows a decline in the number of fish being harvested and a local muskellunge release award program implemented in 1987 has logged over 1000 releases of fish at least 44″ in length. Adult muskellunge monitoring in eleven spawning areas revealed an increase in mean total length of over 63 mm (>2.5 inches) after the regulation changes. Monitoring of age-0 muskellunge by use of seining surveys (1997–2005) indicates consistent reproductive success with the potential for several strong year-classes. Improvements in the muskellunge population and fishery are attributed to the progressive management action and a united community response.     

A program for successful muskellunge management—A Minnesota success story

Minnesota has established a successful muskellunge, Esox masquinongy, stocking program as a result of a series of research studies that enabled area fisheries managers to make informed management decisions. The previous propagation and stocking program (pre-1982) reared muskellunge progeny from Shoepack Lake near the Minnesota–Ontario border, but these fish were not attaining trophy size. Protein electrophoresis showed there were two different muskellunge strains in the state. A controlled study was conducted on progeny from natural muskellunge populations from Leech and Shoepack lakes. Fish from these populations, together, were used to stock two other Minnesota lakes, and their growth rates, age of maturity, and maximum size attained were compared. Leech Lake muskellunge grew faster and attained a larger maximum size than the Shoepack Lake strain. As a result, a new stocking program based on the Leech Lake strain was developed. Because it was difficult to collect Leech Lake gametes, the Minnesota Department of Natural Resources created seven brood stock lakes using Leech Lake progeny. As a result of combining genetic considerations, new research on spawning sites, more restrictive harvest regulations, and catch-and-release fishing, anglers’ catch of trophy-sized muskellunge in Minnesota has increased.     

Detection of VHSV IVb within the gonads of Great Lakes fish using in situ hybridization

Viral haemorrhagic septicaemia virus (VHSV) genotype IVb was recently detected as the cause of numerous mortality events in Great Lakes fish. In situ hybridization was used to examine the gonads from 13 fish, including freshwater drum Aplodinotus grunniens and muskellunge Esox masquinongy that were infected naturally, as well as rainbow trout Oncorhynchus mykiss and fathead minnows Pimphales promelas, which were experimentally infected. Although the ovaries and testes of fish infected by VHSV IVb had few lesions, viral RNA was present in the ovaries of the rainbow trout and fathead minnow and was abundant in the gonads of muskellunge and in the ovaries of freshwater drum. Viral RNA was present mainly surrounding yolk vacuoles/granules or adjacent to the germinal vesicle, with lesser amounts found within the germinal vesicle, in the mesovarium and/or tunica albuginea and blood vessels of the ovary. Viral RNA was also found in and surrounding primary and secondary spermatocytes of the muskellunge.     

Fluctuations in the muskellunge (Esox masquinongy Mitchill) population of Chautauqua Lake,New York

Synopsis Population and exploitation estimates were made from angler recaptures of Chautauqua Lake muskellunge,Esox masquinongy Mitchill. Fish were tagged during Conservation Department studies in 1941–1946, 1961–1965 and 1976–1978. Population estimates of adult fish ranged from one to seven fish per hectare and angler exploitation rates of tagged fish fluctuated from 3.8% to 14.1%. Relative catch indicators suggest a major decline in the lake's muskellunge population during the last decade. Overexploitation, habitat alteration and interspecific competition with recently introduced fish species were cited as probable causes of the decline.     

Evaluation of temperature-selection differences among juvenile muskellunge originating from different latitudes

Genetic differences among muskellunge Esox masquinongy populations are related to residence in major river drainages, suggesting the existence of divergent stocks. By use of radio-telemetry we compared different seasonal and diel temperature selection in a southern Illinois reservoir for three geographically and genetically distinct stocks of age-2 muskellunge from throughout the latitudinal range of the species. Muskellunge from the Upper Mississippi River drainage were represented by the Leech Lake, Minnesota, population and the Ohio River drainage was represented by the Cave Run Lake, Kentucky, population. Progeny from North Spring Lake, Illinois, an interstock, or mixed-origin stock, were also evaluated. No differences in temperature selection were observed among stocks of juvenile muskellunge across seasons or diel periods. The seasonal mean temperatures of the water varied significantly—spring 21.7°C, summer 28.4°C, and fall 14.8°C, with an overall average temperature of 24.1°C, in agreement with previously published values obtained in laboratory trials. This lack of different temperature-selection patterns among stocks is in contrast with other life-history characteristics, for example growth, metabolism, and conversion efficiency, which have been shown to differ among populations and stocks.     

The New River, Virginia, muskellunge fishery: population dynamics, harvest regulation modeling, and angler attitudes

Although muskellunge, Esox masquinongy, fisheries in northern US states and Canadian provinces are increasingly being managed by introduction of restrictive harvest regulations (e.g. 1370-mm (54′′) minimum length limits), many southern US muskellunge fisheries continue to be managed with comparatively liberal regulations (e.g. 762-mm (30′′) minimum length limits) that are implemented statewide. We studied the population dynamics of the New River, Virginia, muskellunge fishery and used predictive modeling to determine whether restrictive harvest regulations also might prove beneficial for this southern latitude fishery. A creel survey was also conducted to learn more about angler attitudes to the New River muskellunge fishery. Muskellunge grew quickly, with fish reaching harvestable lengths (762 mm, 30′′) in 2–3 years. Muskellunge fishing pressure, harvest rates, and voluntary release rates were low compared with reports for more northern areas. Most anglers, irrespective of how often they fished for muskellunge, defined “trophy” muskellunge to be approximately 1050–1100 mm (41–43′′) in length. Although angler support for restrictive harvest regulations was low, abundance of memorable-length (≥1070 mm, 42′′) muskellunge was predicted to increase under all evaluated length limits. Muskellunge yield would remain static at 914-mm (36′′) and 1016-mm (40′′) length limits, because of the rapid growth of fish, but yield would decline dramatically with a 1143-mm (45′′) length limit, because male muskellunge rarely exceeded 1100 mm (43′′). Because of rapid growth and low release rates, implementation of higher length limits (e.g. 965–1067 mm, 38–42′′) may indeed prove beneficial for augmenting “trophy” muskellunge production on the New River. Angler support for higher minimum length limits might be increased by educating anglers about the rapid growth rates of muskellunge and the expected size structure changes that will result from a length-limit increase. Size structure changes resulting from an increase in the minimum length limit may be difficult to detect because of potential increases in fishing pressure or reduced fish growth as a result of competition for food resources. Long-term monitoring of muskellunge growth and angling pressure may therefore be needed to ensure that new regulations are indeed benefitting the fishery.     

Variation in routine metabolism of juvenile muskellunge: evidence for seasonal metabolic compensation in fishes

Metabolic rate of age 0 muskellunge Esox masquinongy ranged from 0·10 at 5° C to 0·24 mg O₂ g^-1 h^-1 at 25° C and was significantly higher in spring and autumn than during winter months at comparable water temperatures. Reduced metabolic rate in winter was consistent with the metabolic compensation hypothesis, implying that metabolism of muskellunge varies independently of acclimation temperature and gonadogenesis. Moreover, seasonal variation in metabolic rate has important implications for energy budget studies. Single-season estimates of esocid metabolism may be inadequate to describe annual energy requirements; the magnitude of errors will depend on the time of year metabolic rate was measured. As a result, it is suggested that seasonal variation in metabolic rate be incorporated into energy budget determinations for fishes.     

Potential effects of spawning habitat changes on the segregation of northern pike (<Emphasis Type="Italic">Esox lucius</Emphasis>) and muskellunge (<Emphasis Type="Italic">E</Emphasis>. <Emphasis Type="Italic">masquinongy</Emphasis>) in the Upper St. Lawrence River

Changes in spawning habitat of northern pike (Esox lucius) may affect their segregation from and coexistence with the closely related muskellunge (E. masquinongy). We estimated the areal coverage of robust and shallow emergent vegetation in three shared-spawning bays in the Upper St. Lawrence River from aerial photographs taken from 1948 to 2003. Robust emergent vegetation (e.g., cattail) increased in coverage by 155–241% while shallow emergents (sedges) decreased by 46–96%. The loss of sedges, an important northern pike-spawning habitat, may facilitate greater spawning overlap in offshore-submersed aquatic vegetation within bay habitats used by muskellunge. Development rates and characteristics of northern pike and muskellunge eggs and larvae were compared to better understand the implications of greater spawning overlap. Northern pike eggs developed faster than muskellunge eggs at temperatures of 4.7–19°C, and adhesive eggs and the presence of adhesive papillae were present in both species. Equations were used to predict degree-day requirements for hatching and swim-up in three habitats (shallow emergents, bay, and offshore shoal) along a temperature gradient. Northern pike required more estimated degree days to reach hatching in bay and offshore shoal habitat relative to shallow emergent habitat due to cooler temperatures. Significant spawning overlap is known to occur within bay habitats, but poor success of northern pike in deep bay habitats and overall reductions in abundance are hypothesized to currently buffer muskellunge from potential negative interactions between these species. Guest editors: J. M. Farrell, C. Skov, M. Mingelbier, T. Margenau & J. E. Cooper International Pike Symposium: Merging Knowledge of Ecology, Biology, and Management for a Circumpolar Species     

Determining minimum ultimate size, setting size limits, and developing trophy standards and indices of comparable size for maintaining quality muskellunge (Esox masquinongy) populations and sports fisheries

Growth and ultimate size can provide important population insights and a sound biological basis for setting length limits, which can be the best single regulation for preventing overexploitation of muskellunge (Esox masquinongy) populations. A system was developed, using cleithral age and total length at age confidence limits (CL) data, to determine reproductive and growth potential (ultimate size) for calculating and setting increased size limits based on minimum reproductive size (upper 99% CL at age at first maturity + 2 year) and minimum ultimate size (MUS) calculated from the lower 99% CL—minimum ultimate size limit (MUSL). MUS also provides a trophy standard and an index of relative size for comparing trophy potential of individuals within and among populations. Guidelines are provided for determining minimum sample size (mean ± 95% confidence interval = 12 ± 4) and minimum age (8–10 ± 2.0 year) required to produce valid von Bertalanffy growth trajectories. MUS, MUSL, and trophy standards for both length and estimated weight are provided for female and male muskellunge from 14 Ontario sources. Mean MUS, or trophy standard, for females was 115 ± 10.3 cm (MUSL range 75–135) and 11.1 ± 2.6 kg (2.5–17.5) and for males was 95 ± 7.5 cm (66–110) and 6.1 ± 1.3 kg (1.9–9.2). These indices can precisely define growth and growth potential for muskellunge populations and individuals and can be used to better manage and maintain or improve the quality of muskellunge populations and fisheries. Dedicated to the late Dr. E.J. Crossman.