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Population genetics of Muskellunge in the St. Lawrence River, its main tributaries and inland lakes of Québec

Quentin Rougemont1, Anne Carrier2, Jeremy Le-luyer3, Anne-Laure Ferchaud1, John M. Farrell4, Daniel Hatin5, Philippe Brodeur6, Louis Bernatchez1

1Département de biologie, Institut de biologie intégrative et des systèmes (IBIS), Université Laval, G1V 0A6, Québec, Canada
2Département de techniques du milieu naturel, Centre d’études collégiales à Chibougamau, Cégep de Saint-Félicien, Chibougamau, G8P 2E9, Canada
3IFREMER, Unité Ressources Marines en Polynésie, Centre Océanologique du Pacifique – Vairao – BP 49 – 98179 Taravao – Tahiti – Polynésie Française
4Department of Environmental and Forest Biology, State University of New York, College of Environmental Science and Forestry, 13210, Syracuse, New York, USA.
5Ministère des Forêts, de la Faune et des Parcs, Direction de la gestion de la faune de l’Estrie-Montréal-Montérégie-Laval, 201, Place Charles-Le Moyne, Longueuil, Québec, J4K 2T5, Canada
6Ministère des Forêts, de la Faune et des Parcs, Direction de la gestion de la faune de la Mauricie et du Centre-du-Québec, 100, rue Laviolette, bureau 207, Trois-Rivières, Québec, G9A 5S9, Canada

Introduction

Over the past decades, an increasing number of fish species have undergone strong decrease in their abundance due to various human activities. Such activities may prevent the free movement of fish, generates pollution and habitat loss, overfishing and many additional problems. To overcome these demographic declines, numerous stocking programs have been implemented to sustain fish populations worldwide. This is the case of the Muskellunge (Esox masquinongy) in the province of Québec, Canada. The species is renowned for his trophy-size specimens which are highly prized by anglers. However, Muskellunge has undergone strong decline in abundance during the first half of the 20th century in the waters of the St. Lawrence River, especially in the greater Montréal region. Consequently, Muskellunge from Ontario and New York State were used for stocking over 1.5 million of individuals from 1950 to 1997. From 1950 to 1965, eggs initially taken from the Chautauqua Lake (New York State, USA) were transferred to the Lachine government hatchery in Québec where fry were reared before being released into the St. Lawrence River, several of its main tributaries and inland lakes. From 1965 to 1986, adults from Lake Joseph were used as source for stocking. Finally, from 1986 to 1997, eggs from Lake Tremblant were used. Muskellunge populations from Joseph and Tremblant Lakes were originally introduced with fish from the Lake Chautauqua source (see details about stocking history in Carrier et al.).

An optimal management of Muskellunge can only be achieved through a detailed understanding of its population structure and of the extant of connectivity between distinct populations. In particular, the existence of genetically differentiated populations with some level of reproductive isolation must be taken into account for sound conservation and management practices. Moreover, genetically distinct groups of fish may evolve to local adaptation in response to particular habitat characteristics (temperature, water chemistry, etc.). Those adaptations allow fish from distinct populations to optimize their reproduction and survival in a given habitat. It is therefore necessary to preserve the natural genetic variation present within a species in order to ensure its potential to evolve and survive in an ever changing environment. Such knowledge will be fundamental to define management units for fishery management, habitat protection and restoration, which is particularly important in highly connected systems such as the St. Lawrence River and its tributaries. Finally, genetics can inform on the extent of hybridization that may have occurred between wild, local fish and fish artificially introduced by stocking.

The genetic structure and diversity of Muskellunge within the St. Lawrence River, its major tributaries and inland lakes of Québec have never been studied. Therefore, the present study has been realised to : 1) evaluate the level of genetic structure in Muskellunge, 2) measure the impact of historical stocking on the genetic structure and diversity, and 3) define evolutionary significant units relevant for population management and to maintain a sustainable resource for angling.

Sampling and genetic characteristics

A total of 662 Muskellunge have been captured in 22 sites for approximately 24 fish per location (Figure 1). Those samples were essentially obtained with the help of professional fishing guides (Mr. Marc Thorpe, Mr. Mike Lazarus, and Mr. Michael Philips), their customers, volunteer sport fishermen, and wildlife technicians. A tiny portion of a pelvic fin was clipped for each fish (1 cm², 100 mg) and preserved in alcohol for further genetic analyses in the laboratory of L. Bernatchez at Laval University in Québec. All fish were released after capture.

This extensive sampling made it possible to cover the sections of the St. Lawrence River, from the Thousand Islands region to Lake Saint-Pierre, its major tributaries and some inland lakes of Québec. The major sources of stocking were also sampled : 1) Chautauqua (New York State) and Pigeon (Kawartha Lakes system, Ontario) Lakes, 2) Joseph Lake, and 3) Tremblant Lake. Muskellunge was introduced in these last two lakes and were used as sources for stocking a few years later. Finally, Lake Traverse located in the Mauricie region of Québec has never been stocked to our knowledge and was included in our study.

DNA from individual fish was extracted from the preserved biopsies in the laboratory. This DNA was then characterized using a new sequencing technology allowing to read each DNA variation over a large part of the fish genome. It was then possible to identify for each fish, over 16 000 genetic variants. Such variants were compared among individuals and among sampling sites, which allowed quantifying the genetic diversity of the species, its population structure and document the impact of past stocking events on the genetic makeup of wild populations.

Figure 1 - Location of sampling sites.
Figure 1 – Location of sampling sites.

Population genetic structure

The genetic results revealed a moderate level of genetic diversity compared with other freshwater fish species that have been studied using similar methods. The effective size of populations, estimated from genetic data, is the number of broods that reproduce effectively, thus transmitting their genetic background to their offspring. In general, the total number of fish in a population can be 10 to 100 times higher than the number of effective individuals. Effective population sizes were generally quite low among Muskellunge populations, especially for isolated lakes. In the St. Lawrence River, the estimate of the effective size was 669 for all sites grouped together. This value is considered moderate compared to other freshwater species, but reflects the unique characteristics of the Muskellunge life-cycle (high longevity, highest position in the food chain, solitary and territorial behaviours) and its typically low population density. These findings highlight the vulnerability of this species and the importance of applying specific protection measures to ensure its sustainability.

The measures of genetic differentiation and population structure suggest the existence of eight distinct genetic groups in the system under study. The first group includes the Muskellunge used as stocking sources and the sites directly derived from this source, namely Chautauqua, Joseph, Tremblant, Frontière, and Maskinongé Lakes, as well as the Chaudière and Saint-Maurice Rivers. This confirms the common origin of the muskellunge of these water bodies, all derived from the source of Lake Chautauqua (New York State). For the Chaudière and Saint-Maurice Rivers, available knowledge suggest that Muskellunge abundance was initially low in those systems, and that stocking would have established perennial populations. The second group corresponds to the l’Achigan River and the third group to the Yamaska River, which are genetically distinct from the St. Lawrence River. The fourth group consists of all sites within the St. Lawrence River, from the Thousand Islands to Lake Saint-Pierre. The fifth group is Lake des Deux-Montagnes, which is also genetically distinct from the Muskellunge of the whole St. Lawrence River. It is noteworthy that Muskellunge from Lake des Deux-Montagnes show a certain proportion of migration to Lake Saint-Louis. Most of these migratory individuals (83%) were found on the North shore of Lake Saint-Louis, which is fed by water coming from the Ottawa River. The sixth group consists of isolated lakes that have never been stocked, represented in this study by Lake Traverse. This body of water has a unique genetic makeup that needs to be preserved. The seventh group corresponds to Pigeon Lake (Kawartha Lakes system in Ontario), used for stocking to a lesser extent than other water bodies, and the eighth group is Lake Champlain.

Although the St. Lawrence River formed a single population, the genetic differentiation between individuals increased with distance between them. This pattern is a consequence of the geographically reduced dispersal of individuals across the entire St. Lawrence River. In addition, the extant of genetic variation observed in the St. Lawrence River proves to be continuous, that is, there are no real, highly differentiated genetic groups. This suggests that dispersal can occur freely from upstream to downstream, although it is obviously reduced upstream by the presence of the two major obstacles on the St. Lawrence River (Beauharnois and Moses-Saunders dams).

Figure 2 - Histogram showing the percentage of each individual belonging to the different genetic groups. Each vertical bar corresponds to an individual sampled in a given body of water and represents its degree of belonging (or mixture) to a given group. Each color represents a genetically distinct group. For example, there is considerable genetic similarity between individuals in Frontière Lake, Joseph Lake and Tremblant Lake (FRO, JOS, and TRE respectively), all of which were seeded from the Chautauqua Lake (CHQ) source. Conversely, there is great genetic distinction between Lake Traverse Muskellunge (TRA) and all other bodies of water. Orange dots : source of individuals used for sowing. Green dots : lakes and rivers where the muskellunge was absent or in low abundance before stocking. Blue dots : sections of the St. Lawrence River and des Deux-Montagnes Lake. For abbreviations meaning, see Figure 1.
Figure 2 – Histogram showing the percentage of each individual belonging to the different genetic groups. Each vertical bar corresponds to an individual sampled in a given body of water and represents its degree of belonging (or mixture) to a given group. Each color represents a genetically distinct group. For example, there is considerable genetic similarity between individuals in Frontière Lake, Joseph Lake and Tremblant Lake (FRO, JOS, and TRE respectively), all of which were seeded from the Chautauqua Lake (CHQ) source. Conversely, there is great genetic distinction between Lake Traverse Muskellunge (TRA) and all other bodies of water. Orange dots : source of individuals used for sowing. Green dots : lakes and rivers where the muskellunge was absent or in low abundance before stocking. Blue dots : sections of the St. Lawrence River and des Deux-Montagnes Lake. For abbreviations meaning, see Figure 1.

Stocking effects

Fine scale analysis of genetic mixing patterns allowed us to estimate the effect of stocking on the genetic structure of populations (Figure 2). This analysis revealed that stocking had very little effect on the genetic integrity of wild populations in the St. Lawrence River. Indeed, we found very little evidence of genetic mixing of Chatauqua, Joseph or Tremblant Lakes strains used as source populations. Conversely, there is evidence of pronounced genetic mixing in some lakes and tributaries of the St. Lawrence, despite the fact that they have in most cases received smaller quantities of stocked fish than the St. Lawrence River. This is the case for the Saint-Maurice and Chaudière Rivers, as well as for Maskinongé Lake, where there was a mixture of local (represented in black in Figure 2) and introduced (represented in green in Figure 2) genetic makeups. The main hypothesis likely to explain this pattern is that stocking has had variable effects depending on the initial size of the populations being stocked. In general, it is expected that stocking done with individuals from different genetic groups, in this case individuals from distant lakes (differences in climate and habitat types), is potentially ineffective due to lack of adaptation of stocked individuals to local conditions. It is therefore possible that the individuals stocked in the St. Lawrence River had a low reproductive success and/or that hybrids resulting from reproduction were poorly adapted to local conditions, ultimately showing a low survival rate. Thus, non-native Muskellunge may have been displaced in the St. Lawrence, which potentially had a larger population size than isolated lakes or tributaries.

Management implications

Our results suggest that from a genetic point of view, the entire St. Lawrence River, from the Thousand Islands region to Lake Saint-Pierre, can be considered as a single population within which genetic differentiation of individuals increase slightly with distance. Thus, a single management unit would be sufficient on the St. Lawrence River to ensure the maintenance of genetic diversity in this system. Of course, individuals who are isolated by impassable obstacles should be managed locally. This is particularly the case of Lake St-François, enclosed by dams upstream (Moses-Saunders) and downstream (Beauharnois). The second management unit includes Lake des Deux-Montagnes, which is genetically different from the St. Lawrence River population. The third group consists of the tributaries of the St. Lawrence River, each representing a distinct unit with some nuance depending on the abundance of Muskellunge prior to stocking. Thus, l’Achigan and Yamaska Rivers showed little evidence of hybridization with stocked fish while the Chaudière and Saint-Maurice Rivers have a more pronounced genetic mixing profile with the stocking sources. The fourth group consists of lakes stocked directly from Chautauqua Lake (Joseph, Tremblant, and Frontière Lakes) which all share a strong genetic similarity with Chautauqua Lake. The fifth group includes lakes into which the Muskellunge was initially present (Maskinongé and Champlain Lakes) where apparently only modest mixing occurred. Finally, Traverse Lake is one of the few, if not the only unstocked natural population in Québec with a unique genetic makeup.

In conclusion, in systems previously unoccupied by Muskellunge or with a very low density of individuals, stocking has made it possible to sustain local populations in the long term and therefore, have helped to enhance recreational fishing activities. Although stocking has temporarily contributed to the species recruitment and to the Muskellunge fishery in the Montréal region of the St. Lawrence system (see the article of Carrier et al. in the present issue), it does not appear to have been successful in the long term, possibly because of the poor adaptation of the stocked individuals to the particular local conditions of a large river such as the St. Lawrence. However, they may have contributed to sustain the fishery on the short term (see Carrier et al. in the present issue). During your next fishing trip, for example on the St. Lawrence River or on Lake des Deux-Montagnes, you will be able to assert that you most likely caught native Muskellunge of local origin. Based on the results of this study, we recommend avoiding future stocking without detailed knowledge of stock abundance, diversity and genetic structure, and of the level of exchange between them. Actions aiming habitat protection and restoration should rather be prioritized in order to optimize the success of natural reproduction.

Acknowledgements

We express our gratitude to muskies anglers who collected most of the samples, especially to Marc Thorpe, Mike Lazarus and Michael Phillips. We thank Christopher Legard (New York State Department of Environmental Conservation) for collecting and sharing Chautauqua Lake samples. We thank Samuel Cartier for collecting Lake Champlain fish. Thanks to Chris Wilson (Aquatic Research and Monitoring Section, Ontario Ministry of Natural Resources and Forestry) for sharing Pigeon Lake DNA samples and to Christopher Wilson (Fish Culture Section, Ontario Ministry of Natural Resources and Forestry) for sharing hatcheries and stocking history between Ontario and Québec. Thanks to Shawn Good (Vermont Fish & Wildlife Department) and Jeffrey J. Loukmas (New York State Department of Environmental Conservation) for sharing historical management and stocking history of Lake Champlain. We also thank Nicolas Auclair, Florent Archambault, Rémi Bacon, Christian Beaudoin, Anabel Carrier, Chantal Côté, Julie Deschesnes, François Girard, Guillaume Lemieux, Louise Nadon, Yves Paradis, Geneviève Richard, and Éliane Valiquette for logistic, field and laboratory assistance. Funding was provided by the ministère des Forêts, de la Faune et des Parcs du Québec, Canadian Research Chair in Genomics and Conservation of Aquatic Resources, Fondation héritage faune (Fédération québécoise des chasseurs et pêcheurs), Ressources Aquatiques Québec, and Muskies Inc.

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Identification of essential Muskellunge habitats in Lake Saint-Pierre

Photo : MFFP

Introduction

This project is part of an initiative undertaken since 2010 by the ministère de la Faune et des Parcs (MFFP) and its many partners to update knowledge about Québec Muskellunge and optimize its management. To measure the current state of the Muskellunge sport fishery in the St. Lawrence River and the des Deux-Montagnes Lake, a survey of the catches was conducted from 2010 to 2013, with the collaboration of three professional fishing guides. In the section of the river located between Montréal and Sorel and in Lake Saint-Pierre, the lower abundance of young specimens harvested by sport fishermen suggested lower recruitment of young Muskellunge in these two bodies of water, compared to Lakes Saint-Louis and des Deux-Montagnes (see Carrier et al. in the present issue for more details).

Some anthropogenic activities have a negative impact on the St. Lawrence ecosystem. They have recently resulted in a deterioration of aquatic habitats, particularly in Lake Saint-Pierre. Nearly 5 000 ha of fish breeding, nursery, and growth habitats in the floodplain has been altered due to the intensification of agricultural practices over the last three decades (de la Chenelière et al. 2014). The loss of large areas of submerged aquatic vegetation beds since the mid-2000s (Figure 1; Magnan et al. 2017) and the proliferation of benthic cyanobacteria (Hudon et al. 2012), which develop on the bottom of Lake Saint-Pierre, have also been documented. These wetlands represent growth habitats and refuges for several fish species. This situation raises fundamental questions about the potential effects of habitat loss on a large predator species such as Muskellunge. In addition, critical breeding and growth habitats of juvenile Muskellunge have never been identified in Lake Saint-Pierre, which limits our ability to properly protect and restore habitats of this species. A study was therefore initiated to monitor adult Muskellunge movements during the spawning and growth seasons, to determine habitat characteristics selected by fish and to locate breeding and rearing areas of juveniles.

Figure 1 - Abundance of submerged aquatic vegetation in Lake Saint-Pierre from 2002 to 2016 (from Magnan et al. 2017).
Figure 1 – Abundance of submerged aquatic vegetation in Lake Saint-Pierre from 2002 to 2016 (from Magnan et al. 2017).

Methodology

Figure 2 - Radio (bottom picture) and acoustic (top picture) transmitters used for Muskellunge marking. Credit : MFFP.
Figure 2 – Radio (bottom picture) and acoustic (top picture) transmitters used for Muskellunge marking. Credit : MFFP.

The identification of movement patterns and the precise location of Muskellunge has been made possible through the use of advanced telemetry technologies.

Two types of transmitters have been inserted into the fish abdomen: an acoustic transmitter, which is detected by stationary receivers at strategic locations in the St. Lawrence River and its tributaries (Figure 2) and a radio transmitter including an external antenna visible on the ventral portion of the fish, allowing precise location of the specimens using a mobile receiver, operated from a plain or a boat (Figure 3). In spring 2017 and spring 2018, a total of about 80 fixed receivers were deposited annually near the waterbed between Montréal and the Gentilly sector (Figure 4). These receivers, recovered at the end of each autumn, continuously record the passage of tagged fish. The number of the individual, the date and time of passage are then extracted and used for migration analysis purposes.

Figure 3 - Mobile radio receiver (left picture) and fixed acoustic receiver (right picture) used to locate Muskellunge. Credit : MFFP.
Figure 3 – Mobile radio receiver (left picture) and fixed acoustic receiver (right picture) used to locate Muskellunge. Credit : MFFP.

This information provides information on habitat use and residence time of Muskellunge in various sections of the St. Lawrence River and its tributaries. They also help defining the periods and patterns of seasonal migration of the species. In addition, accurate, real-time fish telemetry locations provide information on the location and characteristics of adult staging sites during spring breeding and during summer and fall growth seasons.

Figure 4 - Location of fixed acoustic receptors used to measure the passage of Muskellunge tagged in Lake Saint-Pierre in 2018. The stations installed between Gentilly and Québec city, to the right of this map, were not represented.
Figure 4 – Location of fixed acoustic receptors used to measure the passage of Muskellunge tagged in Lake Saint-Pierre in 2018. The stations installed between Gentilly and Québec city, to the right of this map, were not represented.

Preliminary results

A total of 21 Muskellunge were caught by sport fishing thanks to the valuable collaboration of two professional anglers, Mr. Mike Lazarus and Mr. Marc Thorpe, and to the MFFP wildlife technicians. The fish were surgically fitted with transmitters during the fall of 2016 and fall of 2017. Females and males ranged from 38 to 52 inches in size (Figure 5). The implementation of the transmitters carried out by the wildlife technicians of the MFFP went very well. All Muskellunge were located on at least one occasion, approximately 6 to 18 months after being tagged, indicating that all individuals survived after surgery.

Ten individuals tagged in the fall of 2016 were followed by boat and plain between April 25 and May 24, 2017. During this period, which includes migration to breeding sites and spawning activities, 112 locations of individual Muskellunge were noted. The habitat selected by each individual was also characterized (vegetation, substrate, temperature, current velocity, oxygen concentration, depth, etc.). The locations recorded in the spring of 2017 showed that all Muskellunge tagged at Lake Saint-Pierre during the previous fall used the Lake Saint-Pierre area to reproduce. The data revealed that 38 % of radio tagged individuals used Lake Saint-Pierre tributaries during the breeding season (April-May). Specimens were located in the du Loup, Saint-François and Nicolet Rivers, as well as in the Chenal Tardif (a section of the Saint-François River). After breeding, these individuals migrated to feeding habitats in the St. Lawrence River. The rest of the individuals used Lake St. Pierre wetlands during the spawning season. In the spring, Muskellunge were found at depths ranging from 0.6 to 8.2 m (mean : 3.1 m), in low current velocity, mostly lower than 0.1 m/s. In the majority of cases, Muskellunge were found in habitats showing submerged vegetation of moderate to high abundance.

The analysis of movements recorded in 2017, based on data collected by dozens of fixed receivers, showed that after the breeding season, the majority of fish tagged in the fall of 2016 in Lake Saint-Pierre spent some time in this area during the summer and fall of 2017. However, during summer, 60% of the individuals made large-scale migrations towards the stretch of the river located between Montréal and Sorel. Some Muskellunge even reached the stations located near the Jacques Cartier Bridge in Montréal.

In order to track the movements of the 21 tagged individuals, the telemetry monitoring work will continue in 2018 and 2019. All the results collected during this project will enable to identify and map the preferential habitats for Muskellunge, particularly for reproduction, which could be protected or restored as needed. The preliminary results of 2017 already underline the role of the shallow marshes of Lake Saint-Pierre and some of its tributaries for the reproduction of the species. It will be important to validate these observations over the next few years, to estimate the contribution of these various sectors to the recruitment of the species and to evaluate the state of health of habitats. In addition, the long-distance migrations reported in 2017 emphasize that the management of Muskellunge and its habitats must be done at the scale of the entire studied fluvial section, including the downstream portion of the tributaries. This finding is supported by the results of the genetic structure of populations, which demonstrated the homogeneity of the genetic signature of the Muskellunge population in the St. Lawrence stretch located between Lake Saint-Louis and Lake Saint-Pierre (see Rougemont et al. in the present issue for details on population genetics).

Figure 5 - Length frequency distribution of marked Muskellunge in Lake Saint-Pierre.
Figure 5 – Length frequency distribution of marked Muskellunge in Lake Saint-Pierre.

Warning to anglers

If you catch a marked Muskellunge, you must release it after noting the fish and telephone number written on the tag that is inserted at the base of the dorsal fin (it is often necessary to scrape the surface of the label to clearly see the numbers therein). Be careful, it is important to avoid taking the specimen out of the water and to limit the handling time. Moreover, these tips apply to all Muskellunge catches, whether tagged or not. Then contact a MFFP biologist at the number written on the tag to provide the date, location of capture, and, if possible, pictures of the ventral portion of the fish.

Acknowledgements

We would like to thank all the partners who participated in the financing and the realization of the telemetry work. Special thanks to professional fishermen Mike Lazarus and Marc Thorpe for their support throughout the development of the study and for their participation in the capture of the specimens. Thanks also to Florent Archambault, Nicolas Auclair, Rémi Bacon, Virginie Boivin, Chantal Côté, Charles-Étienne Gagnon, Guillaume Lemieux, Yves Paradis, and René Perreault for their support and for all efforts in the field. The project is made possible by the collaboration and financial support of the ministère des Forêts, de la Faune et des Parcs, of the Comité ZIP du lac Saint-Pierre, of Muskies Canada, of the Fondation de la faune du Québec, of Thomas marine, of the Fondation héritage faune (Fédération québécoise des chasseurs et des pêcheurs) and of some private donors.

 

References

De la Chenelière, V., P. Brodeur et M. Mingelbier (2014). Restauration des habitats du lac Saint-Pierre : un prérequis au rétablissement de la perchaude. Le Naturaliste canadien. 138 (2) : 50-61.

Hudon, C., A. Cattaneo, A.-M. Tourville Poirier, P. Brodeur, P. Dumont, Y. Mailhot, Y.-P. Amyot, S.-P. Despatie and Y. De Lafontaine (2012). Oligotrophication from wetland epuration alters the riverine trophic network and carrying capacity for fish. Aquatic Sciences. 74 : 495-511.

Magnan, P., P. Brodeur, É. Paquin, N. Vachon, Y. Paradis, P. Dumont et Y. Mailhot (2017). État du stock de perchaudes du lac Saint-Pierre en 2016. Comité scientifique sur la gestion de la perchaude du lac Saint-Pierre. Chaire de recherche du Canada en écologie des eaux douces, Université du Québec à Trois-Rivières et ministère des Forêts, de la Faune et des Parcs. vii + 34 pages + annexes.

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Québec Muskellunge : Two centuries of fishing and management history

Gustave Provost, directeur de la station piscicole de Lachine en 1962. Gustave Prévost, director of the Muskellunge hatchery in 1962. Crédit : MFFP.

Anne Carrier ¹ ², Philippe Brodeur³, Daniel Hatin⁴ and Louis Bernatchez¹
¹Département de biologie, Institut de Biologie Intégrative et des Systèmes (IBIS), Université Laval, G1V 0A6, Québec, Canada
²Département de Techniques du milieu naturel, Centre d’études collégiales à Chibougamau, Cégep de Saint-Félicien, Chibougamau, G8P 2E9, Canada
³Ministère des Forêts, de la Faune et des Parcs, Direction de la gestion de la faune de la Mauricie et du Centre-du-Québec, 100, rue Laviolette, bureau 207, Trois-Rivières, G9A 5S9, Canada
⁴Ministère des Forêts, de la Faune et des Parcs, Direction de la gestion de la faune de l’Estrie-Montréal- Montérégie-Laval, 201, Place Charles-LeMoyne, Longueuil, Québec, J4K 2T5, Canada

Muskellunge is one of the most mythical and impressive fish species. Over the past two centuries, biologists and Muskellunge anglers have documented many fascinating aspects of its biology. For example, it’s impressive size (Bernatchez and Giroux 2012), its unusual migration abilities (Kerr and Jones 2017) and even its surprising reproductive behavior (Crossman 1990, Jennings et al. 2011). The history regarding Muskellunge is fascinating, as evidenced by the origin of its name and the history of its management, which reveal the particular importance of Muskellunge in Québec.

This article is a non-exhaustive historical overview of some of the most important aspects of Muskellunge management in Québec. It includes some historical references regarding the nomenclature and taxonomy of Muskellunge, its original and contemporary spatial distribution and the stocking history. This article reports the work done as part of a Master’s thesis, which first aimed at gathering available historical information that would support the interpretation of genetic data on Muskellunge in Québec waters (see article of Rougemont et al. in this issue).

Taxonomy and Québec folklore

As early as the colonization time of New France, documents from the Société Provancher mentions that the first viceroy of France, Sieur Jean-François La Rocque de Roberval, used the basin of the Maskinongé River as his fishing territory. At the time, Muskellunge was a well-known species as evidenced by the multiple presumed Amerindian roots of its name, which meant big pike, ugly pike or spotted pike (Crossman 1986, MacCaughey 1917). Gradually, these appellations have derived to become « long mask » or « elongated mask » in Québec French. Today, the two generally accepted names are « maskinongé » in Canada and « Muskellunge » in the United States, but there are between 40 and 94 common names in French only (see Mellen 1917, Chambers 1923, Weed 1927 and Crossman 1986 for an exhaustive inventory of the different names and their origin). As mentioned by Crossman (1986), probably no other fish has, in a single language, as many forms or spelling of its common name. According to Weed (1927), the number of its common names is a fairly reliable index of the extent to which a fish attracts attention. This partly explains this diverse nomenclature, but as Mongeau (1976) points out, this taxonomic confusion also certainly comes from its great resemblance to the Northern pike (Esox lucius) and the fact that it has been recognized quite lately as a different species from his cousin.

Commercial fishing and natural distribution in the 19th century

Since the nomenclature of the species was highly variable until the beginning of the 20th century, it is very difficult to interpret observations regarding the Muskellunge distribution until the 1900s. In the 19th century, Muskellunge was highly prised by native and non-native anglers and, because of the quality of its flesh and its imposing size, it contributed to a significant commercial fishery in Québec. Although today the opinions are mixed about the taste of the Muskellunge flesh, the naturalist Constantine Rafinesque mentioned in 1818 that « it is one of the best fish (…) its flesh is very delicate and divides easily like salmon, in large white patches like snow » (MacCaughey 1917). According to historical records of the Canadian fisheries management authorities (Crossman 1986), nearly 2.9 million pounds, representing approximately 192 535 Muskellunge, were harvested by the commercial fishery in Québec from 1868 to 1936. Interestingly, commercial catches of Muskellunge in the waters of the Montréal area accounted for 90 % of the landings of this species throughout the province (Fry et al. 1942). Muskellunge commercial fishing ceased in 1936.

The historical texts suggest that native Muskellunge was found only in southern Québec, even if its northern and southern distribution limits are only very slightly defined. Its distribution was likely limited to the waters of the St. Lawrence River watershed and some of its tributaries from the Ottawa River to Québec City (Small 1883, Dymond 1939, Vézina 1977). According to information available at the end of the 19th century, native Muskellunge was found from the southern border of the province (including the Champlain Lake and the Richelieu River watersheds) to Northwest of Outaouais, Laurentides, Lanaudière, and Mauricie regions (Dymond 1939). Specifically, Dymond (1939), Small (1883), Halkett (1906 and 1907), and Montpetit (1897) report that Muskellunge was present (1) in the Rideau River north of Merrickville (Outaouais, Québec), (2) in the Ottawa River south of Rapides des Joachim (MRC de Pontiac, Outaouais, Québec), south of the Petawawa River and up to Travers Lake (Algonquin Provincial Park, Ontario), and (3) in several lakes connected to the Gatineau and du Lièvre Rivers, including Gilmour, Donaldson, and Plumbago Lakes (MRC Collines-de-l’Outaouais, Outaouais, Québec). In addition, some isolated populations were discovered in 1968 after the dismantling of private fishing clubs in the Mauricie region, specifically in the des Envies River watershed, which is a tributary of the Batiscan River, where the Traverse Lake (Potvin 1973, Pageau et al. 1978) analyzed in the study of Rougemont et al. (see article in this issue) is located. Finally, according to the interpretation of Fry et al. (1942), quoted by Robitaille and Cotton (1992), the most important native population in Québec would have been in Lake St-Louis, a fluvial lake of the St. Lawrence River.

Active management period

Stocking

Gustave Provost, directeur de la station piscicole de Lachine en 1962. Gustave Prévost, director of the Muskellunge hatchery in 1962. Crédit : MFFP.
Gustave Prévost, director of the Muskellunge hatchery in 1962. Credit : MFFP.

Muskellunge has been one of the most stocked fish species in Québec (Dumont 1991). Prior to 1950, few Muskellunge stocking in Québec were recorded in the literature (MacCaughey 1917, Dymond 1939, Small 1883, Halkett 1906 and 1907). At the end of the first half of the 20th century, a significant decline of Muskellunge populations in the waters of the St. Lawrence River and of the Montréal Archipelago, associated with overfishing and habitat loss, raised worries and questions. Therefore, the wildlife management authorities undertook a major restoration project which included the construction of the very first Muskellunge hatchery facility in Lachine (borough of Montréal city, Québec) (Pictures 1 to 3), as well as the development of a local expertise on esocids breeding (Vezina 1977). In 1950, these actions led to the beginning of stocking, which were adapted to contemporary knowledge in 1985. Muskellunge stocking continued until 1997. During the same period, the species was also introduced, with or without success, in more than 80 Québec water bodies in order to create new opportunities and enhance existing Muskellunge populations (Vézina 1977, Dumont 1991, Vincent and Legendre 1974, Brodeur et al. 2013, de la Fontaine, Y. unpublished). In a few rare cases, Muskellunge introduction has been used in an attempt to control competing species in brook trout lakes. Introducing a top-predator into the food chain obviously had an impact on the fish communities.

Photo 2 - Muskellunge hatchery facility in Lachine (1950-1964). Credit : MFFP.
Photo 2 – Muskellunge hatchery facility in Lachine (1950-1964). Credit : MFFP.

Muskellunge farming began in Québec at the Lachine hatchery in 1950. Due to water supply problems, breeding was transferred to the Baldwin Mills hatchery in 1964 (now known as the Baldwin-Coaticook provincial hatchery) (Dumont 1991). Following unsuccessful attempts to breed Muskellunge from several local lakes such as Lake des Deux-Montagnes (Montréal area) and the Gilmour, Donaldson, and Plumbago Lakes (Outaouais) (MPC 1961, Vezina 1977, Crossman and Goodchild 1978), embrocated eggs were imported from the Bemus Point hatchery (New York, USA) and, to a lesser extent, from the Deer Lake hatchery (Ontario, Canada) to start production (Kerr 2001, Dufour and Paulhus 1977, Christopher Wilson and Christopher Legard, personal communication). Muskellunge from both hatcheries originated respectively from the Chautauqua Lake (New York, USA) and from Stony Lake, Buckhorn Lake, and from the Crowe River, these three last water bodies being part of the Kawartha Lakes system in Ontario. According to the information we gathered, it appears that all the lakes used by these hatcheries have also been stocked with an unknown Muskellunge source to support their respective fishery (Christopher Wilson and Christopher Legard, personal communication). Both of these hatcheries, as well as the one of Lachine, are no longer in operation.

Photo 3 - Muskellunge transport from the Lachine hatchery. Credit : MFFP
Photo 3 – Muskellunge transport from the Lachine hatchery. Credit : MFFP

From 1965 to 1986, Joseph Lake (Centre-du-Québec, Québec) was used as a broodstock source to supply the Baldwin Mills hatchery (Dumont 1991). Subsequently, from 1986 to 1997, Lake Tremblant (Laurentides, Québec) was used as the source population. Muskellunge was originally introduced in both lakes from the American or Ontarian sources (see Figure 1 – simplified stocking history in Québec). The results of the genetic study confirmed that the American source was the most likely for both lakes.

Stocking, carried out over several decades in the Montréal area, has been effective in improving the stock status and maintaining the Muskellunge sport fishery. In fact, an analysis of Muskellunge recruitment measured from 1962 to 1977 revealed that 55 % of the annual abundance of young Muskellunge could be explained by the number of yearly stocked individuals and the abundance of young Muskellunge stocked the previous year (cannibalism
and/or competition effects) (Dumont 1991). In 1998, the improvement of the Muskellunge population structure, distributed over a long time period, and the presence of natural recruitment justified the end of stocking (Cloutier 1987, Dumont 1991). Since then, no Muskellunge stocking has been done in Québec.

Figure 1 - Simplified representation of stocking in the St. Lawrence River and some inland lakes of Québec. Arrows represent stocking events from the different source populations. Full arrows show clear mentions of stocking, while the dotted arrows reflect anecdotal mentions.
Figure 1 – Simplified representation of stocking in the St. Lawrence River and some inland lakes of Québec. Arrows represent stocking events from the different source populations. Full arrows show clear mentions of stocking, while the dotted arrows reflect anecdotal mentions.

Integrating collaborative science to Muskellunge management

In parallel to the management actions undertaken by the Québec government, a general reflection on fishing practices and a growing interest in the conservation of a high quality fishery focusing on trophy-size specimens emerged, leading to the creation of Muskies Canada (Wachelka 2008a,b,c) and to the beginning of a long collaboration between muskies anglers and the Québec wildlife management authorities. Muskellunge is not vulnerable to capture by the scientific fishing gears used to monitor fish communities in the St. Lawrence River. Monitoring the sport harvest of Muskellunge through angling surveys is therefore an excellent alternative to contribute to its management and to allow evaluation of the effectiveness of the management measures.

To evaluate the status of Muskellunge stocks, a study was conducted in the 1990s in collaboration with the Montréal chapter of Muskies Canada. From 1994 to 1997, five anglers tagged and released 808 Muskellunge, mainly in the Montréal area. The results showed that a few hours of fishing were enough to catch a Muskellunge, whereas in the 1970s, an experienced angler needed approximately 100 hours of fishing to catch a single specimen. After three years of survey, 88 tagged fish were recaptured by anglers, which corresponded to a recapture rate of 11 %, considered relatively low and indicative of a total Muskellunge abundance of several thousands of specimens (Pierre Dumont, personal communication) The gradual increase in the extent of the Muskellunge size structure suggested by the fishing surveys and the presence of a natural production of young Muskellunge justified the cessation of stocking in 1998 (Dumont 1991).

To update the data on the Muskellunge fishery in the St. Lawrence River (from Lake Saint-François to Lake Saint-Pierre) and in Lake des Deux-Montagnes, a second survey was conducted from 2010 to 2013, more than a decade after stocking ended. This second study was conducted with the invaluable collaboration of three professional anglers recognized in Québec, Mr. Marc Thorpe, Mr. Mike Lazarus and Mr. Michael Phillips. A total of 2 569 Muskellunge were captured, of which 2 162 were tagged by three volunteer anglers. Of these tagged fish, 108 were recaptured. The order of magnitude of recapture rates was low in all studied sectors (3.7 % to 4.8 %). Compared to the study carried out in the Montréal area from 1994 to 1997, the recapture rate reported from 2010 to 2013 was twice lower (4.8 % compared to 11 %). Since the recapture rate is generally inversely proportional to the total abundance of a population, this result suggests that the abundance of Muskellunge in the Montréal area has increased since the stocking ended, at least for medium to high size fish, targeted by anglers.

According to archived data from 1918 to 1927, 19 % of Muskellunge caught in Lake Saint-Louis exceeded the legal minimum size of 44 inches (Figure 2). In 1973, this proportion was of 16 % and then increased to almost 50 % in the late 1990s and to 54 % during the 2010-2013 period. This improvement over several decades can be explained by stocking, combined with the enforcement of a minimum legal size of 38 inches in 1986, which has been increased to 44 inches in 1998 (Figure 2). Because of the presence of large specimens, the waters of the St. Lawrence River and of Lake des Deux-Montagnes are now identified as sites of great interest for Muskellunge anglers. In the section of the St. Lawrence River between Montréal and Lake Saint-Pierre, the low abundance of young specimens smaller than 35 inches in the Muskellunge sport harvest suggests a lower recruitment, compared to Lake Saint-Louis and Lake des Deux-Montagnes (Figure 3). This result justified the realization of a study conducted by the ministère des Forêts, de la Faune et des Parcs (MFFP) and its numerous partners that aims to identify the essential habitats of the species by using telemetry
(see the article of Brodeur et al. in this issue).

Figure 2 - Historical comparisons of the proportion of fish larger than 44 inches caught by sport fishing on Lake St-Louis. The year of introduction of minimum sizes to 38 inches in 1986, increased to 44 inches in 1998, is also represented.
Figure 2 – Historical comparisons of the proportion of fish larger than 44 inches caught by sport fishing on Lake St-Louis. The year of introduction of minimum sizes to 38 inches in 1986, increased to 44 inches in 1998, is also represented.

The most recent fishery survey has generated some preliminary knowledge about Muskellunge migration. Thus, between 2010 and 2013, the majority of marked individuals (95 %) recaptured by the sport fishery within six months after tagging or one to two years after, were in the same body of water where they had been tagged. The distances measured between specimens capture and recapture were generally less than a few kilometers, both on a one year scale and between years (72.7 % and 58.1 % of recaptures within 5 km from the tagging location, respectively). This result suggests that, although Muskellunge can travel long distances, particularly during the breeding season, a large proportion of individuals return to specific areas corresponding generally to large vegetation beds favorable to feeding. This result demonstrates the importance of preserving and restoring the submerged aquatic vegetation beds of the St. Lawrence River. However, large-scale movements between the various sectors of the river have been observed between Lake Saint-Pierre and the Montréal-Sorel section, with distances of up to 58 km. This result was recently corroborated by the preliminary results of the telemetry study, which shows that a certain proportion of the Muskellunge tagged at Lake Saint-Pierre migrate upstream during the feeding season (see article by Brodeur et al. in this issue). These observations of large scale movements also corroborate the connectivity existing throughout the St. Lawrence River system revealed by genetic analyses.

Figure 3 - Size structure of Muskellunge caught by sport fishing during the 2010- 2013 period in the St. Lawrence River watersheds (LDM: Lake des DeuxMontagnes, LSF: Lake Saint-François, LSL: Lake Saint-Louis, MS: stretch between Montréal and Sorel, LSP: Lake Saint-Pierre). The proportion of fish greater than or equal to 44 inches, 36 to 43 inches and 35 inches or less is shown.
Figure 3 – Size structure of Muskellunge caught by sport fishing during the 2010- 2013 period in the St. Lawrence River watersheds (LDM: Lake des Deux-Montagnes, LSF: Lake Saint-François, LSL: Lake Saint-Louis, MS: stretch between Montréal and Sorel, LSP: Lake Saint-Pierre). The proportion of fish greater than or equal to 44 inches, 36 to 43 inches and 35 inches or less is shown.

Future perspectives

To maintain the trophy status of the species, which can maintain and improve the quality of the Muskellunge fishery, a regular review of the stock status and management is required. Since 2010, a study aiming at gathering new knowledge on several aspects of Muskellunge biology has been conducted by the MFFP and its numerous partners. This vast study will contribute to Muskellunge management in Québec. To date, this initiative has led to a retrospective of historical management, reported in this article, to a genetic analysis of Muskellunge populations (see article by Rougemont et al. in this issue), and to a study aiming to identify essential Muskellunge habitats between Montréal and Lake Saint-Pierre. Some anglers report a recent decline in the quality of the Muskellunge fishery in some inland water bodies of Québec, which remains to be measured. Muskellunge studies based on angling surveys have thus been underway for some years in the Maskinongé Lake and the Ottawa River (see Deschesnes in this issue).

Acknowledgements

We thank the following people for their valuable collaboration. We would like to acknowledge the involvement of the Muskellunge anglers who participated to the 2010-2013 angling survey: Marc Thorpe, Mike Lazarus, and Michael Phillips. Special thanks to Peter Levick (Muskies Canada), Chris Wilson (Aquatic Research and Monitoring Section, Ontario Ministry of Natural Resources and Forestry), and John Farrell (Department of Environmental and Forest Biology, State University of New York) who shared with us many information on Muskellunge management. Thanks to Christopher Legard (New York State Department of Environmental Conservation) and Christopher Wilson (Fish Culture Section, Ontario Ministry of Natural Resources and Forestry) for sharing the Chautauqua Lake and Deer Lake hatcheries history. Thanks to Steven Kerr (retired biologist, Fisheries Section, Ontario Ministry of Natural Resources) for his invaluable advice and for sharing his knowledge on the history of Muskellunge management in Québec. Thanks to Shawn Good (Vermont Fish and Wildlife Department) and Jeffrey J. Loukmas (New York State Department of Environmental Conservation) for sharing management and stocking history of Champlain Lake. We also thank the Fédération québécoise des chasseurs et des pêcheurs, Ressources Aquatiques Québec and Muskies Inc. for their financial support. Funding was also provided by the ministère des Forêts, de la Faune et des Parcs du Québec and by the Canada Research Chair in Genomics and Aquatic Resources Conservation.

Références

Bernatchez, L. et M. Giroux (2012). Les poissons d’eau douce du Québec et leur répartition dans l’est du Canada. 2e éd., Ottawa, Canada.

Brodeur, P., D. Hatin et R. Bacon (2013). Suivi du maskinongé dans le Saint-Laurent et le lac des Deux-Montagnes. Dans: Atelier sur la faune aquatique, 19-21 février 2013, Sainte-Foy, Québec.

Chambers, E.T.D. (1923). The maskinonge: a question of priority in nomenclature. Transactions of the American Fisheries Society (1922) 52: 171-177.

Cloutier, L. (1987). Le maskinongé (Esox masquinongy). Dans : Problématique de la conservation et de la mise en valeur d’espèces de poissons d’eau douce au Québec. Ministère du Loisir, de la Chasse et de la Pêche, Québec.

Crossman, E. J. (1986). The noble muskellunge: a review. In : Managing muskies: a treatise on the biology and propagation of Muskellunge in North America (éd. Gordon HE), p.1-13. American Fisheries Society, Bethesda, Md.

Crossman, E. J. (1990). Reproductive homing in Muskellunge, Esox masquinongy. Canadian Journal of Fisheries and Aquatic Sciences, 47(9): 1803-1812. doi:10.1139/f90-205

Crossman, E. J. and C. D. Goodchild (1978). An annotated bibliography of the muskellunge, Esox masquinongy (Osteichthyes: Salmoniformes).  https://www.biodiversitylibrary.org/item/123600#page/3/mode/1up

De la Fontaine, Y. (non publié). Muskellunge stocking in southern Québec waters.

Dufour, M. and P. J. Paulhus (1977). L’élevage et l’ensemencement du maskinongé au Québec. Dans : Compte rendu du 10e atelier sur les poissons d’eau chaude, p. 117-127. Ministère du Loisir, de la Chasse et de la Pêche.

Dumont, P. (1991). Les ensemencements de maskinongé, de truite brune et de truite arc-en-ciel dans les eaux de la plaine de Montréal. Dans : Colloque sur l’ensemencement, p. 30-41. Conseil de l’aquaculture et des pêches.

Dymond, J. R. (1939). The fishes of the Ottawa region [version électronique]. https://www.biodiversitylibrary.org/item/111705#page/7/mode/1up

Fry, F., J.-P. Cuerrier et G. Préfontaine (1942). Première croissance du maskinongé dans le lac Saint-Louis en 1941. Dans : Rapport de la Station biologique de Montréal et de la Station biologique du Parc des Laurentides pour l’année 1941, p. 170-175. Fascicule 2, appendice VII, Manuscrit.

Halkett, A. (1906). Report of the Canadian Fisheries Museum. In : 38th Annual report, p. 362-370. Department of marine & fisheries, Fisheries Branch. Appendix number 14.

Halkett, A. (1907). Report of the Canadian Fisheries Museum. In : 40th Annual report, p. 321-349. Department of marine & fisheries, Fisheries Branch. Appendix number 14.

Jennings, M. J., G. R. Hatzenbeler and J. M. Kampa (2011). Spring capture site fidelity of adult muskellunge in inland lakes. North American Journal of Fisheries Management, 31(3): 461-467.

Kerr S. J. and T. A. Lasenby (2001). Esocid stocking: an annotated bibliography and literature review. Fish and Wildlife Branch, Ontario Ministry of Natural Resources. Peterborough, Ontario. 138 p. and appendix.

Kerr J. S. and Jones B. (2017). Movements of Muskellunge in the Saint-John River based on a volunteer tagging project, 2006-2015. American Fisheries Society Symposium, 85: 39-50.

MacCaughey V. (1917). The Chautauqua Mascalonge or Muskalunge. Dans : B. W. Huebsh (dir.), The natural history of Chautauqua (p. 80-83) [En ligne], [https://www.biodiversitylibrary.org/item/71239#page/7/mode/1up].

Miller L. M., J. M. Farrell, K. L. Kapuscinski, K. Scribner, B. L. Sloss, K. Turnquist and C. C. Wilson (2017). A review of muskellunge population genetics: implications for management and future research needs. American Fisheries Society Symposium, 85: 385-414.

Ministère des Pêcheries et de la Chasse de la Province de Québec (1961). Contribution de la station piscicole de Lachine à l’étude de maskinongé. Dans : Journal de bord de l’office de biologie.

Mellen, I.M. (1917). Twenty four ways of spelling the name of a fish (muskellunge). New York Zoology Society Bulletin 20, p. 1558.

Montpetit, A.-N. (1897). Le maskinongé. Dans : Les poissons d’eau douce du Canada, p. 76-80, https://www.biodiversitylibrary.org/item/45738#page/3/mode/1up

Mongeau, J .R. et G. Massé (1976). Les poissons de la région de Montréal, la pêche sportive et commerciale, les ensemencements, les frayères, la contamination par le mercure et les PCB. Ministère du Loisir de la Chasse et de la Pêche, Service de l’aménagement et de l’exploitation de la faune, Montréal, Québec. Rapport technique no 06-13. xviii + 286 p.

Mongeau, J. R., J. Leclerc et J. Brisebois J. (1980). La répartition géographique des poissons, les ensemencements, la pêche sportive et commerciale, les frayères et la bathymétrie du fleuve Saint-Laurent dans le bassin de Laprairie et les rapides de Lachine. Ministère du Loisir, de la Chasse et de la Pêche, Service de l’aménagement et de l’exploitation de la faune. Rapport technique no 06-29. 145 p.

Pageau, G., Y. Gravel and V. Legendre (1978). Distribution and value of the esocidae in Québec waters. Dans : Compte rendu du 10e atelier sur les poissons d’eau chaude, p. 1-7. Ministère du Loisir de la Chasse et de la Pêche, Direction de la recherche faunique.

Potvin, C. (1973). Inventaire ichtyologique du bassin de la rivière des Envies. Découverte de populations indigènes de maskinongé. Ministère du Loisir de la Chasse et de la Pêche, Direction de la recherche faunique.

Robitaille, J. A. et F. Cotton (1992). Bilan des connaissances sur le maskinongé (Esox masquinongy) et sur ses populations dans le Saint-Laurent. Ministère du Loisir, de la Chasse et de la Pêche, Direction de la gestion des espèces et des habitats. Rapport technique, p. 1-55.

Small, H. B. (1883). Fishes of the Ottawa District. Transactions of the Ottawa Field-Naturalists’ Club (1882-1883), 4: 31-49.

Turnquist, K. N., W. A. Larson, J. M. Farrell, P. A. Hanchin, K .L., Kapuscinski, L. M. Miller, K. T. Scribner, C .C., Wilson and B. L. Sloss (2017). Genetic structure of muskellunge in the Great Lakes region and the effects of supplementation on genetic integrity of wild populations. Journal of Great Lakes Research, 43(6): 1141-1152. doi:10.1016/j.jglr.2017.09.005

Vézina, R. (1977). Les introductions de maskinongé, Esox masquinongy, au Québec et leurs résultats. Dans : Compte rendu du 10e atelier sur les poissons d’eau chaude, p. 129-135. Ministère du Tourisme, de la Chasse et de la Pêche du Québec, Service de l’aménagement de la faune.

Vincent, B. et V. Legendre (1974). Répartition géographique du maskinongé, Esox maskinongy, dans le district des Laurentides. Compilation 1972. District de Montréal, Service de l’aménagement de la faune et Service de la recherche biologique. Ministère du Tourisme, de la Chasse et de la Pêche du Québec, Service de l’aménagement de la faune. Rapport technique.

Wachelka, H. (2008a). Muskies Canada, the first 10 Years. Muskies Canada Release Journal, mai/juin, p. 11.

Wachelka, H. (2008b). Muskies Canada, the Middle Years. Muskies Canada Release Journal, juillet/août, p. 11.

Wachelka, H. (2008c). Muskies Canada, 1999 to Present. Muskies Canada Release Journal, septembre/octobre, p. 8-10.

Weed, A. C. (1927). Pike pickered and muskalonge, Zoology leaflet 9. In: D. C. Davies (dir.), Field museum of natural history Chicago, p. 152-205, https://www.biodiversitylibrary.org/item/25559#page/75/mode/1up