The Introduction and Spread of Aquatic Invasive Species ... · Cette publication hautement spécialisée, La propagation d’espèces . aquatiques envahissantes par l’utilisation

The Introduction and Spread of Aquatic Invasive Species through the Recreational Use of BaitA LITERATURE REVIEW JUNE 2014

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The Spread of Aquatic Invasive Species through the Recreational Use of Bait: A Literature Review

Abstract

This document was prepared to summarize information related to the introduction and spread of aquatic invasive species in Ontario through the recreational use of live bait.

Literature on the recreational use of bait as a pathway for the spread of aquatic invasive species is examined. Various pathways are reviewed and evaluated. These pathways include accidental release, escape from holding containers, fellow travellers in holding water and packaging materials, illegal dumping of bait and the commercial sale/use of aquatic invasive species.

Impacts of aquatic invasive species may be direct or indirect. Potential impacts are many and varied including interspecific competition, preda-tion, transmission of disease and parasites, habitat alteration, genetic impacts, displacement of native species, community alteration, and adverse effects on recreational and commercial fisheries.

The use of live bait provides a pathway for the introduction and spread of aquatic invasive species which can potentially result in significant ecological and economic impacts.

© Queen’s Printer for Ontario, 2014Printed in Ontario, Canada Report Prepared by Steven J. Kerr for Biodiversity Policy SectionBiodiversity BranchOntario Ministry of Natural Resources

Suggested citation: Kerr, S. J. 2014. The Introduction and Spread of Aquatic Invasive Species through the Recreational Use of Bait: A Literature Review. Report prepared for Biodiversity Branch.

Ontario Ministry of Natural Resources. Peterborough, Ontario. Ontario Ministry of Natural Resources300 Water StreetPeterborough, ONK9J 8M5

Cover photo: Iowa Darter (Etheostoma excile). Photo: Al Dextrase

La propagation d’espèces aquatiques envahissantes par l’utilisation d’appâts à des fins récréatives : examen des travaux publiés

Sommaire

Le présent document a été rédigé pour présenter le résumé des renseignements sur l’introduction, en Ontario, d’espèces aquatiques envahissantes par l’utilisation d’appâts à des fins récréatives.

Y sont examinés les travaux publiés sur l’utilisation d’appâts à des fins récréatives. Diverses voies d’introduction y sont examinées et évaluées, dont celles-ci : les appâts libérés accidentellement dans des plans d’eau; les appâts qui s’échappent de leur vivier; la présence d’espèces commensales dans les viviers et les matériaux d’emballage; le rejet illégal d’appâts; la vente ou l’utilisation à des fins commerciales d’espèces aquatiques envahissantes.

Les espèces aquatiques peuvent avoir des effets directs et indirects. Les effets possibles sont nombreux et variés. En sont des exemples la compétition interspécifique, la prédation, la transmission de maladies et de parasites, l’altération des habitats, les effets de nature génétique, le déplacement d’espèces indigènes, l’altération des communautés et les effets indésirables sur les pêches récréatives et commerciales.

L’utilisation d’appâts vifs favorise l’introduction et la propagation d’espèces aquatiques envahissantes. Cela peut se répercuter de façon importante sur les écosystèmes et l’économie.

Cette publication hautement spécialisée, La propagation d’espèces aquatiques envahissantes par l’utilisation d’appâts à des fins récréatives : examen des travaux publiés n’est disponible qu’en anglais en vertu du Règlement 671/92 qui en exempte l’application de la Loi sur les services en français.

Pour obtenir de l’aide en français, veuillez communiquer avec le ministère des Richesses naturelles au 1-800-667-1940.

1The Introduction and Spread of Aquatic Invasive Species through the Recreational Use of Bait: A Literature Review

Table of Contents

Common and Scientific Names of Aquatic Species Cited in the Text .........................................................2

Introduction ..................................................................................................................................................3

Pathways for the Transfer and Movement of Aquatic Invasive Species .......................................................4

Ontario’s Bait Industry ..................................................................................................................................5

Recreational Use of Bait as a Pathway for Aquatic Invasive Species ...........................................................6

Accidental Release ................................................................................................................................7

Escape from Holding Containers ..........................................................................................................7

Fellow Travellers in Holding Water and Packaging Materials ..............................................................7

Illegal Dumping of Bait at End of Fishing Trip ......................................................................................7

Commercial Sale and Use of Illegal Aquatic Invasive Species ..............................................................8

Other Pathways .....................................................................................................................................8

Overall Pathway Considerations ...........................................................................................................8

Potential Impacts of Releases of Aquatic Invasive Species ..........................................................................9

Interspecific Competition ......................................................................................................................9

Predation ...............................................................................................................................................9

Transmission of Disease and Parasites ...............................................................................................10

Habitat Alteration ................................................................................................................................10

Genetic Impacts ..................................................................................................................................11

Community Alteration ........................................................................................................................12

Economic Impacts ...............................................................................................................................12

Displacement of Indigenous Species .................................................................................................12

Adverse Effects on Commercial and Recreational Fisheries ..............................................................13

Acknowledgements ....................................................................................................................................14

References and Literature Cited ................................................................................................................14

Personal Communications ..........................................................................................................................25

Websites .....................................................................................................................................................25

Additional References ................................................................................................................................25

General ................................................................................................................................................25

Vectors and Pathways of Aquatic Invasive Species .............................................................................28

Ecological Impacts of Introductions of Non-indigenous Species .......................................................29

Glossary ......................................................................................................................................................36

Appendix 1. Some of the more common potential pathways for the transfer and movement of aquatic invasive species in Ontario. .........................................................37

Appendix 2. Introduction or spread of aquatic invasive species attributed to the recreational use of bait. .............................................................................39

Appendix 3. Impacts of the introduction of aquatic invasive species and pathogens in a new waterbody (modified from Kerr and Grant 2000). ..................................................44

2 The Introduction and Spread of Aquatic Invasive Species through the Recreational Use of Bait: A Literature Review

Common and Scientific Names of Aquatic Species Cited in the Text

AMPHIBIANSCalifornia Tiger Salamander

(Abystoma californiense)

Chiracahua Leopard Frog (Rana chiricahuensis)

Northern Leopard Frog (Lithobates pipiens)

CRUSTACEANSRusty Crayfish (Orconectes rusticus)

Signal Crayfish (Pacifastacus leniusculus)

FISHAmerican (rainbow) Smelt (Osmerus mordax)

Amistad Gambusia (Gambusia amistadensis)

Arkansas River Shiner (Notropis girardi)

Arroyo Chub (Gila orcuttii)

Banded Darter (Etheosoma zonata)

Bandfin Shiner (Luxilus zonistius)

Banff Longnose Dace

(Rhinichthys cataractae smithi)

Black Bullhead (Ameiurus melas)

Blackchin Shiner (Notropis heterodon)

Blacknose Dace (Rhinichthys atratulus)

Blacknose Shiner (Notropis heterolepis)

Blackside Darter (Percina maculata)

Bluegill (Lepomis macrochirus)

Bluehead Sucker (Catostomus discobolus)

Bluntnose Minnow (Pimephales notatus)

Brassy Minnow (Hybognathus hankinsoni)

Brook Stickleback (Culaea inconstans)

Brook Trout (Salvelinus fontinalis)

Brown Hoplo (Hoplosternum littorale)

Bull Chub (Nocomis raneyi)

Bullhead Minnow (Pimephales vigilax)

Central Mudminnow (Umbra limi)

Central Stoneroller (Campostoma anomalum)

Channel Catfish (Ictalurus punctatus)

Colorado Pikeminnow (Ptychocheilus lucius)

Commanche Spring Pupfish

(Cyprinodon elegans)

Common Carp (Cyprinus carpio)

Common Shiner (Luxilus cornutus)

Creek Chub (Semotilus atromaculatus)

Crescent Shiner (Luxilus cerasinus)

Cutlip Minnow (Exoglossum maxillingua)

Delta Smelt (Hypomesus transpacificus)

Emerald Shiner (Notropis atherinoides)

Fallfish (Semotilus corporalis)

Fantail Darter (Etheostoma flabellare)

Fathead Minnow (Pimephales promelas)

Finescale Dace (Chrosomus neogaeus)

Flathead Catfish (Pylodictis olivaris)

Fourspine Stickleback (Apeltes quadracus)

Golden Shiner (Notemigonus crysoleucas)

Goldfish (Carassius auratus auratus)

Hornyhead Chub (Nocomis biguttatus)

Humpback Chub (Gila cypha)

Iowa Darter (Etheostoma exile)

Johnny Darter (Etheostoma nigrum)

Lake Chub (Couesius plumbeus)

Lake Herring (Coregonus artedii)

Largemouth Bass (Micropterus salmoides)

Least Darter (Etheostoma microperca)

Leon Spring Pupfish (Cyprinodon bovinus)

Logperch (Percina caprodes)

Longnose Dace (Rhinichthys cataractae)

Longnose Sucker (Catostomus catostomus)

Lost River Sucker (Deltistes luxatus)

Mimic Shiner (Notropis volucellus)

Mosquitofish (Gambusia affinis)

Mottled Sculpin (Cottus bairdii)

Mountain Redbelly Dace (Chrosomus oreas)

Mountain Sucker (Catostomus platyrhynchus)

Ninespine Stickleback (Pungitius pungitius)

Northern Hog Sucker (Hypentelium nigricans)

Northern Pearl Dace (Margariscus margarita)

Northern Pikeminnow

(Ptychocheilus oregonensis)

Northern Redbelly Dace (Chrosomus eos)

Orangethroat Darter (Etheostoma spectabile)

Oriental Weatherfish

(Misgurnus anguillicaudatus)

Owens Sucker (Catostomus fumeiventris)

Pecos Pupfish (Cyprinodon pecosensis)

Plains Minnow (Hybognathus placitus)

Rock Bass (Ambloplites rupestris)

Round Goby (Neogobius melanostomus)

Rainbow Darter (Etheostoma caeruleum)

Rainbow Shiner (Notropis chrosomus)

Razorback Sucker (Xyrauchen texanus)

Red Shiner (Cyprinella lutrensis)

Redfin Shiner (Lythrurus umbratilis)

Red River Shiner (Notropis bairdi)

Rio Grande Silvery Minnow

(Hybognathus amarus)

River Chub (Nocomis micropogon)

River Darter (Percina shumardi)

Rosefin Shiner (Lythrurus ardens)

Rosyface Shiner (Notropis rubellus)

Round Goby (Neogobius melanostomus)

Rudd (Scardinius erythropthalmus)

Ruffe (Gymnocephalus cernua)

Sacramento Sucker ( Catostomus occidentalis)

San Marcos Gambusia (Gambusi georgei)

Sand Shiner (Notropis stramineus)

Santa Ana Sucker (Catostomus santaanae)

Sea Lamprey (Petromyzon marinus)

Sheepshead Minnow (Cyprinodon variegatus)

Shorthead Redhorse

(Moxostoma macrolepidotum)

Shortnose Sucker (Chasmistes brevirostris)

Shoshone Sculpin (Cottus greenei)

Silver Redhorse (Moxostoma anisurum)

Slimy Sculpin (Cottus cognatus)

Southern Redbelly Dace

(Chrosomus erythrogaster)

Speckled Dace (Rhinichthys osculus)

Spottail shiner (Notropis hudsonius)

Spotfin Shiner (Cyprinella spiloptera)

Striped Shiner (Luxilus chrysocephalus)

Tench (Tinca tinca)

Tessellated Darter (Etheostoma olmstedi)

Threespine Stickleback (Gasterosteus aculeatus)

Tonguetied Minnow (Exoglossum laurae)

Trout-Perch (Percopsis omiscomaycus)

Tui Chub (Siphateles bicolor)

Utah Chub (Gila atraria)

Wakasagi (Hypomesus nipponensis)

Walleye (Sander vitreus)

White Perch (Morone americana)

White Shiner (Luxilus albeolus)

White Sucker (Catostomus commersonii)

Yellowfin Shiner (Notropis lutipinnis)

Yellow Perch (Perca flavescens)

MOLLUSCSScrew Snail (Thiara tuberculata)

Zebra Mussel (Dreissena polymorpha)

ZOOPLANKTONSpiny Water Flea (Bythotrephes longimanus)


The introduction and spread of invasive species and pathogens is a global issue (Mills et al. 1994, Lodge et al. 2000a, Lintermans 2004, Chandra and Gerhardt

2008). Introductions of non-indigenous species is considered one of the greatest threats to global biodiversity second only to direct habitat destruction (Convention of Biological Diversity 2000). The rate of introduction of aquatic species has increased steadily over the past two hundred years coincident with increases in global trade (Welcomme 1988, Ricciardi 2006, Hulme 2009) and a variety of human-mediated mechanisms (Hulme 2009). Ricciardi (2006) reported the current human-mediated rate of invasion of aquatic species was 1.8 species per year (i.e., one new species every 28 weeks) which has led to cumulative increases in the diversity of non-indigenous taxa in many aquatic ecosystems. For example, there are currently more than 180 non-indigenous aquatic organisms in the Great Lakes alone (Eyraud and MacIsaac 2013).

While not all species moved by humans become established in a new environment, many can and do. The success or failue of establishment is generally determined based on the ecological filters concept (Kolar and Lodge 2002) (see Glossary). This concept assumes that patterns of establishment are based on characteristics of the species being transported and the suitability of recipient ecosystems. As a result, establishment rates will vary from area to area. For a number of invasive plants and animals in Great Britain Williamson and Fitter (1996) concluded that one of ten imported species would become established in the wild. Conversely, in Europe, the average rate of establishment of an invasive species was estimated to be 63% (Garcia-Berthou et al. 2005). In some cases (e.g., introduction of marine organisms into a freshwater environment), the establishment rate can be effectively zero. Once an alien species becomes established, eradication is seldom possible or practical because of the spatial scale of impacted environments and the general lack of suitable eradication mechanisms (Mack et al. 2000). Thus many resource management agencies seek methods to reduce the means by which invasive species may be introduced or spread. Clearly, it is most important to prevent undesired introductions from occurring.

The economic impacts of aquatic invasive species are large and significant. For example, Leigh (1998) estimated that the costs of implementing an eleven year Ruffe control program in Lake Superior would cost $12 million. Costs of Zebra Mussel removal and control in the United States have been estimated at more than $100 million per year (Pimentel et al. 2000). In Ontario, Zebra Mussels are estimated to cost $75-100 million annually (Marbek Inc. 2010). Annual costs of monitoring and controlling the invasive Sea Lamprey in the Great Lakes

Introduction

is approximately $13 million (Jenkins 2001). Lovell and Stone (2005) estimated that the economic losses associated with invasions of non-indigenous fish in the United States approximated $1 billion annually.

In the future, the effect of the world’s changing climate is expected to enhance the ability of many non-indigenous species to expand their ranges (Hellmann et al. 2008, Rahel and Olden 2008). Mandrak (1989) predicted that at least twenty-seven additional fish species would be capable of invading the Great Lakes basin and have a dramatic impact on resident Great Lakes fish communities. It will be important to preventatively manage pathways for current and future patterns of invasion (Leung et al. 2002).

Inland harvest. Photo: A. Drake 2011


Pathways for the Transfer and Movement of Aquatic Invasive Species

There are a variety of pathways and vectors (see Appendix 1) that allow aquatic invasive species to be transported between donor and recipient ecosystems. In many cases it is impossible to discriminate amongst various pathways and vectors. Many of these pathways are considered to be human-induced dispersal however (Mills et al 1993).

As human activity increases so does the probability of expanded colonization by new species. Increased global trade and mobility of anglers and boaters have increased the risk of spreading aquatic invasive species (Hulme 2009). There is also a geographic scale to consider. Primary pathways, such as international shipping, ballast water and canals, involve movements across broad ecoregions while secondary pathways (e.g., anglers, boaters, etc.) usually involve movements at smaller geographic scales.

Aquatic ecosystems seem to be especially vulnerable to biological invasion (Reaser 2003). Aquatic invasive species can spread more quickly than many terrestrial organisms (Peters and Lodge 2009). Once an invasive species becomes established in freshwater, they often disperse more quickly through interconnected lakes and rivers (Drake 2011).

This document has been prepared to summarize knowledge and information regarding pathways and impacts of aquatic invasive species with particular reference to the commercial sale and use of bait in Ontario’s recreational fishery.

Mimic Shiner (Notropis volucellus). Photo: Al Dextrase


Ontario’s Bait Industry

In Ontario, bait may be harvested for personal use or some types can be purchased from a commercial retailer (see review by Kerr 2012). Species which can be harvested for personal use include crayfish, fish, northern leopard frogs, and leeches. There is no regulatory control over other species, such as earthworms, maggots, hellgrammites, etc., which are also used as bait in Ontario. The commercial bait industry is comprised of both harvesters and dealers which are licenced by the Ontario Ministry of Natural Resources. Species which can be harvested for sale include designated fishes (see Table 1) and leeches. Target fish species are small-bodied fishes in the families Cyprinidae, Percidae, Catostomidae, Gasterosteidae, Salmonidae, Esocidae, Cottidae and Percopsidae (Drake 2011).

Between 2002-2010, it was estimated that the harvest of wild baitfish averaged 4.7 million dozen fish per year (Kerr 2012). Harvest of wild bait occurs throughout the province but the majority of the wild baitfish harvest, for commercial sale, occurs in southern Ontario and the lower Great Lakes (Drake 2011). There is limited culture of the ten baitfish species that are eligible for culture under the regulations of the Fish and Wildlife Conservation Act (Brenda Koenig personal communication).

It has been estimated that there are approximately 4.2 million angling events involving baitfish each year in Ontario (Drake and Mandrak 2014a). This statistic involved any trip where bait was either harvested or purchased. The Emerald Shiner is one of the most commonly used baitfish in Ontario.

In 1980, Ontario’s bait industry was valued at $12.4 million (Can.) (Litvak and Mandrak 1993). Litvak and Mandrak (1993) estimated that the bait industry in North America was valued at $1.06 billion (Can.) annually of which the Ontario bait industry was $30.8 million (Can.) annually. Based on sales, Ontario’s bait industry has more recently been valued at $24.5 million (Can.) (OMNR 2009).

Table 1. Fishes designated as baitfish in Ontario.

MINNOWS

Blacknose Shiner

Blackchin Shiner

Bluntnose Minnow

Brassy Minnow

Central Stoneroller

Common Shiner

Creek Chub

Eastern Blacknose Dace

Emerald Shiner

Fallfish

Fathead Minnow

Finescale Dace

Golden Shiner

Hornyhead Chub

Lake Chub

Longnose Dace

Mimic Shiner

Northern Redbelly Dace

Northern Pearl Dace

Redfin Shiner

River Chub

Rosyface Shiner

Sand Shiner

Spotfin Shiner

Spottail Shiner

Striped Shiner

SCULPINS

Mottled Sculpin

Slimy Sculpin

SUCKERS

Longnose Sucker

Northern Hog Sucker

Shorthead Redhorse

Silver Redhorse

White Sucker

STICKLEBACKS

Brook Stickleback

Ninespine Stickleback

Threespine Stickleback

DARTERS AND LOGPERCH

Blackside Darter

Fantail Darter

Iowa Darter

Johnny Darter

Least Darter

Rainbow Darter

River Darter

Tesselated Darter

Logperch

MISCELLANEOUS

Central Mudminnow

Lake Herring (cisco)

Trout-Perch


Recreational Use of Bait as a Pathway for Aquatic Invasive Species

There are several generic mechanisms which allow bait-related activities to pose a risk of invasion (Andrew Drake personal communication). The first involves the bycatch of non-target (i.e., invasive) species by anglers or commercial harvesters. Secondly, there is the movement potential afforded through commercial harvest and sale, anglers, etc. Finally, the third key variable involves angler behaviour. These variables need to be considered in order to consider invasion risk and evaluate management options (Drake and Mandrak 2014b).

The recreational use of bait is recognized as one of several major pathways for the movement and spread of aquatic invasive species (Drake 2011). On a global scale, Kolar and Lodge (2000) concluded that the global bait trade was increasing and had a strong interaction with aquatic invasive species. Internationally, Welcomme (1988) attributed 11 introductions to the escape or release of bait. Mills et al. (1993) reported that 19% of introduced aquatic invasive species in the Great Lakes basin could be attributed to the recreational use of bait. Upon reviewing literature associated with this exercise, the introduction of almost 130 aquatic species could be attributed to the bait pathway (see Appendix 2).

Many anglers use bait during their angling activities. In North America the use of live bait is largely concentrated in states and provinces in or near to the Great Lakes drainage (Drake 2011, Drake and Mandrak 2014b). Drake and Mandrak (2014b) estimated that approximately 81% of Ontario resident licenced anglers used live baitfish at some point in a given year. This represents an estimated average of eight live baitfish trips/year per individual angler.

Of the anglers that utilize live baitfish in a given year, an estimated 46.7% purchase their bait but not self harvest it, 2.1% will self harvest their bait but not purchase it, and 51.1% will use both methods of baitfish supply (Drake and Mandrak 2014b). Ludwig and Leitch (1996) reported that 57% of anglers in North Dakota and Minnesota, USA, utilized live bait. Most (53%) of these anglers purchased bait from a retailer.

Anglers display substantial overland movement potential and frequently cross drainage basin boundaries to reach angling destination waterbodies (Ludwig and Leitch 1996, Drake and Mandrak 2010, Drake 2013). Anglers may travel large distances from the site they purchase their bait to the water they intend to fish. For example, using a gravity model and variables such as the density of licenced anglers, road travel distance, lake surface area and quality of recreational fishery, Drake and Mandrak (2010) estimated the median distance of aggregated movement was 292.6 km but that lengthier distances were possible (75th percentile = 484.6 km; 95th percentile = 775.2 km. Ludwig (1995) found that a significant

number of surveyed anglers indicated their intent to move bait from the Mississippi River drainage to the Hudson Bay drainage.

With the popularity of bait amongst anglers, there is the potential of transfers and releases of non-indigenous organisms into new waters (commonly referred to as bait bucket transfers – see Glossary). The problem arises when anglers empty the contents of their bait bucket into the water they were fishing instead of destroying the contents on shore. Bait releases accounted for 16.4% of all non-indigenous fishes introduced to inland waters of the United States (Fuller et al. 1999). Kilian and Klauda (2013) attributed the use of live bait for introducing five fish species and four species of crayfish into Maryland waters. Litvak and Mandrak (1999) reported that more than 100 species are believed to have been released into new waterbodies in the United States through bait bucket releases. They concluded that many of the range extensions observed in North America could be attributed to bait bucket transfers. Ludwig and Leitch (1996) concluded that significant measures were required to prevent the dispersal of non-indigenous species by anglers.

The spread of the invasive crayfish (Orconectes rusticus) has been attributed to several vectors including bait bucket releases (Lodge et al. 2000a). Upon review of the potential vectors for non-indigenous crayfish introductions, Lodge et al. (2000b) concluded that the live bait industry was the most important vector for introductions of non-indigenous crayfishes.

Dip netting baitfish. Photo: Brenda Koenig


The use of bait has been identified as the source of many introductions of non-indigenous aquatic species (see Appendix 2). There are six basic means by which aquatic invasive species can be spread via the bait pathway:

ACCIDENTAL RELEASE

Accidental release involves unintentional releases. Some examples could include spilling the contents of a bait bucket while angling, escape of a baitfish off the hook, or highway accidents where contents of holding equipment are spilled. While situations like these undoubtedly occur, it is not believed that this is a major pathway for the introduction or spread of aquatic invasive species.

ESCAPE FROM HOLDING CONTAINERS

Holding containers are used by both anglers and bait harvesters. Anglers may use containers to hold relatively small numbers of bait for their personal use. Bait harvesters often use nets or large containers to offload bait for short term storage. Escapes can occur if there are holes or if an inappropriate mesh size is used. During extreme storm events and flooding, holding containers may be damaged allowing escape of the contents.

FELLOW TRAVELLERS IN HOLDING WATER AND PACKAGING MATERIALS

Non-indigenous organisms can be spread either by introducing the species themselves or by the water or packing materials they are held and transported in. Early life stages of many invaders (e.g., Zebra Mussels, Spiny Water Flea, etc.) are virtually invisible to the naked eye and can be transferred in water held in bait buckets, live wells and boat bilges. Neary and Leach (1992) concluded that bait buckets was the pathway of most significance for the spread of Zebra Mussels to inland waters of Ontario.

Packaging and holding materials may also serve as a vector for transferring other fauna. In a survey of several northeastern Atlantic states, Haska et al. (2012) found that 13 species of macroalgae and 23 species of invertebrates were associated in packing materials of bait worms. Cohen (2012) reported that four microbial species, two species of seaweed, and 30 species of invertebrate were detected in shipments of marine bait shipped from Maine to California.

ILLEGAL DUMPING OF BAIT AT END OF FISHING TRIP

The mobility of anglers allow them to serve as a potential vector for the introduction of species at considerable distances from their native range (Goodwin et al. 2004, Drake 2011, Drake and Mandrak 2014a). Several surveys have revealed that angler release of live bait to the wild is a common activity despite the fact the activity is illegal. In a survey of Ontario anglers, Litvak and Mandrak (1993) reported that 41% of anglers released the contents of their bait bucket at the end of their fishing trip. In another Ontario survey, Dextrase and McKay (1999) found that 46% of anglers released bait after fishing. A survey of Ontario anglers and boaters conducted in 2009 indicated that approximately 36% of respondents released leftover bait (crayfish, frogs, fish and leeches) into the water where they were fishing (LURA Consulting 2010). Kulwicki et al (2003) reported this activity for 36% of anglers in Wisconsin and Michigan. Based on a social survey initiated in 2006, Drake and Mandrak (2014b) estimated that 29.9% of Ontario live bait anglers (either those who purchased or self-harvested their bait) released their leftover fishes at least some of the time although release rates displayed substantial variation across the province (e.g., northern vs. southern Ontario). The release of purchased bait was estimated to occur 893,029 times per year in Ontario (Drake and Mandrak 2014a). Using the same survey, an estimated 9.5% of self-harvesting anglers would release their bait following movement of fishes to another waterbody (Drake and Mandrak 2014b). In Maryland, USA, it was estimated that 65% and 69% of anglers released their unused baitfish and crayfish, respectively (Kilian et al. 2012). In a 2008 survey of Canadian and U.S. jurisdictions, 49% of respondents identified bait bucket introductions as contributing to the spread of alien crayfish species (DiStefano et al. 2009). Koehn (2004) attributed the rapid spread of Carp in Australia to releases by anglers who used Carp as bait.

Releasing unused bait is largely based on two social variables: bait release is convenient and many anglers believe that released fish provide forage for other fishes. The perception of bait release as providing a forage resource is a common theme among anglers who release their bait (Andrew Drake personal communication).

Leeches harvested in Ontario. Photo: Brenda Koenig


COMMERCIAL SALE AND USE OF ILLEGAL AQUATIC INVASIVE SPECIES

Bycatch of non-target species by bait harvesters can occur as evidenced by the occurrence of non-target and invasive fishes in dealers and anglers purchases (Drake 2011). The probability of capturing non-target species during commercial harvest is expected to vary in relation to factors such as the composition of harvest ecosystem, harvest strategy (e.g., site selection, capture technique, etc.) and degree of harvest effort. Based on a generic harvest model involving bait fish in Great Lakes tributary streams (Drake and Mandrak 2014b), the probabililty of capture of the most common non-target species (Rock Bass) (median P = 0.1841) was more than 4,000 times greater compared to the probability of capturing the least likely species (Fourspine Stickleback) (P = 0.00044). The probability of capturing invasive Goldfish and Round Goby as bycatch was moderate (mean P capture values of 0.016 and 0.014, respectively) with these species posing a median 95% chance of capture as bycatch at 186 and 208 harvest events respectively. Therefore, when all else is equal, the probability of bycatch during commercial harvesting increases with harvest effort. Should bycatch exist during commercial harvest the vast majority of these non-target species will be culled by harvesters at the harvest site or elsewhere (Drake and Mandrak 2014b). Dealers contribute further to culling and species sorting so strong overall bycatch reductions from harvest through to sale occur. Drake and Mandrak (2014b) found that there was approximately a 60% reduction for bycatch in the wild compared with the proportion of retail tanks containing non-target species. A 54.6% reduction was observed in the proportion of angler purchases containing bycatch compared with purchases. These results confirm that a strong degree of species sorting is occurring through the pathway. When non-target species are purchased incidentally as bycatch, their abundance is usually quite low although their presence confirms imperfect culling and sorting by harvesters and dealers.

A number of commercial bait shop surveys have documented the presence of prohibited or alien bait species. In a Missouri study, at least 27% of bait shops were found to be selling illegal crayfish species (DiStefano et al. 2009). In a survey of bait shops in Maryland, at least six non-native bait species were documented for sale (Kilian and Klauda 2013). During a 1988 survey of retail baitfish dealers in Toronto, Litvak and Mandrak (1999) found that of the 28 species being held for sale, six species were illegal. In a study of bait retailers in Pennsylvania, LoVullo and Stauffer (1993) reported that 43% of species available for sale were illegal species. Ludwig and Leitch (1996) found that 28.5% of species being held in tanks of Minnesota and North Dakota bait dealers were illegal.

Sales of illegal invasive species can be done knowingly but, in many cases, bait retailers and anglers either don’t know what species are illegal or have difficulty in identifying various species of fish or crayfish. In some cases, specimens of illegal species may be missed during the sorting exercise. In any event, the commercial sale and use of illegal invasive species is known to occur.

Finally, anglers harvesting their own bait may be unable to identify different species and inadvertently contribute to the transfer and spread of aquatic invasive species (Drake and Mandrak 2014b). Some anglers may deliberately use some species of fish (e.g., Gobies, Yellow Perch, Goldfish, etc.) as live bait even though they are illegal.

OTHER PATHWAYS

There are several other potential pathways associated with the recreational use of bait. Sometime the use of dead bait can result in the spread of invasive species. For example, the inadvertent disposal of fertilized eggs from dead American Smelt used as bait has been attributed to the spread of smelt into new waters (Gunderson 2008). In some cases, bait species may be illegally imported from other jurisdictions. This type of activity seems to occur predominantly during the spring walleye fishery and the fall muskellunge fishery (Mark Robbins personal communication).

OVERALL PATHWAY CONSIDERATIONS

Generally, concern for aquatic invasive species associated with the recreational use of bait involves understanding how bycatch, the movement of fishes and risky human behaviour contribute to the introduction and spread of invasive species. In Ontario, the joint occurrence of these activities is relatively rare but, nonetheless, ecologically significant. Given the sheer number of harvest events and live bait angling trips each year, even low cumulative probabilities of bycatch, movement and release of a particular species can be expected to occur at many time and in many places in any one year. Based on an estimated 4.2 million angling trips per year, Drake and Mandrak (2014a) estimated that the most likely number of Round Goby propagules introduced would be 3,715 individuals among 1,288 lakes. Therefore, although bycatch in many bait pathways is relatively low and angler trips with the potential to introduced species are rare, the scale of most fisheries (volume of harvest activity and angling trips) allows these events to occur a sizable number of times in a given year (Ludwig and Leitch 1996).


Potential Impacts of Releases of Aquatic Invasive Species

Impacts of an introduced species can either be direct, by competition or predation, or indirect by causing native species to alter their behaviour (He and Kitchell 1990). There are a number of potential impacts when a non-indigenous species becomes established in a new environment (see review by Goodchild 1999 and Appendix 3).

INTERSPECIFIC COMPETITION

Competition occurs when resources are limited. Trophic interactions can influence habitat choice, survivability and abundance (Link 2002).

Competition for food between native Ninespine Sticklebacks and introduced Threespine Sticklebacks had negative impacts on the native population (Goodchild 1999). Hrabik et al. (1998) found that introduced Rainbow Smelt competed for food with Yellow Perch in Wisconsin lakes. Introduction of the Golden Shiner into California resulted in competition with trout, panfish and Walleye for food (Moyle 1976). This competition for food and space resulted in decreased growth and reproduction in the native species. The Sacramento Sucker, which was introduced to California, was found to compete with native trout for food and space. In addition, their feeding activities disrupted trout redds (Fuller et al. 1999). In New Zealand waters, introduced Rudd competed for invertebrate food resources with native species. Mosquitofish, introduced across the U.S. for mosquito control, had negative impacts on many native small fish populations resulting from competition and predation (Myers 1967, Courtenay and Meffe 1989).

In North Carolina, Feiner et al. (2013) reported that the diet of the invasive White Perch overlapped significantly with several native fish species which resulted in negative impacts to those native fishes. Hurley and Christie (1977) implicated White Perch in the decline in a number of fishes including bluegill, largemouth bass, and walleye in the Bay of Quinte.

Introduced Creek Chub in Québec were found to compete for food with native Brook Trout (Magnan and Fitzgerald 1982). Introduced populations of the Utah Chub were found to become serious competitors with resident sport fish species including trout (Sigler and Miller 1963). It is well known that the introduction of Yellow Perch has served to diminish many Brook Trout fisheries in Ontario and elsewhere (Fraser 1978, Goodchild 1999, Maricle 2007). Competitive interactions between Red Shiners and Fathead Minnows have had an adverse effect on Colorado Pikeminnow (Fuller and Nico 1996).

Non-indigenous crayfish are believed to reduce native populations of crayfish through competition, predation and reproductive interference (Lodge et al. 2000a). In a Wisconsin lake, Lodge et al. (1994) reported strong trophic interactions in which the omnivorous Rusty Crayfish controlled the abundance of macrophytes, snails, and periphyton.

PREDATION

Predation by an invasive species can have a pronounced impact on resident biota particularly when the native stock is already depleted or recovering (Link 2002).

In Lakes Michigan, Huron and Erie, Barbiero and Tuchman (2004) reported a decline in a number of cladoceran species coincident with the invasion of the predatory Spiny Water Flea. They concluded that invasive invertebrate predators could have a pronounced long lasting effect on the zooplankton community. Duxbury et al. (2009) concluded that direct predation after the introduction of Brown Hoplo had an adverse impact on native macroinvertebrate assemblages in Florida waters. Similarly, Marsh and Douglas (1997) concluded that predation from introduced fish could have significant effects, in terms of depleting numbers and recruitment, on native species.

Predation by introduced Rainbow Smelt was responsible for dramatic declines in zooplankton communities in two Wisconsin lakes (Beisner et al. (2003). Post-invasion predation by the North American Signal Crayfish reduced the abundance of two benthic fishes in a British river (Guan and Wiles 1997). Gamradt and Kats (1996) reported that the non-indigenous crayfish and Mosquitofish preyed heavily upon newt larvae. Introduced Red Shiners are known to predate the spawn of the endangered Razorback Sucker (Goodchild 1999). In California, introductions of predatory centrarchids were believed to be responsible for the decline of native frogs as well as the California Tiger Salamander (Dill and Cordone 1997). The Northern Pikeminnow has been noted to prey on several game fish species in areas where it has been introduced (Brown and Moyle 1981).

In addition to direct predation, many invasive small fishes, such as Gobies, also predate the eggs from the nests of resident species (Atkinson 1931, Stenton 1951, Stauffer and Wagner 1979, Fitzsimons et al. 2002). Crayfish are known to consume eggs and larvae of both fish and amphibians (Dorn and Wojdak 2004).


TRANSMISSION OF DISEASE AND PARASITES

Introductions of invasive species can serve to transfer disease and parasites (Stewart 1991).

Lintermans (2004) reported that Oriental Weatherloach, an alien species established in Australia through bait bucket introductions, was host to monogenean parasites which were new to Australia. Kidd (2004) concluded that the use of frogs as bait provided the opportunity to spread the infectious disease ranavirus. Golden Shiners are known to be host for the protozoan parasite Plistophora ovariae (Litvak and Mandrak 1993). LoVullo and Stauffer (1993) reported that cyprinids imported from New York and sold in Pennsylvania were infested with glochidia, a parasite. In a similar event, the introduced European trematode (Bolbophorus confuses) was transferred from South Dakota to Arkansas via a shipment of minnows (Goodchild 1999).

The introduction of Red Shiners to Utah is believed to have been the means by which the Asian tapeworm entered the Virgin River (Fuller et al. 1999). The bass tapeworm (Proteocephalus ambloplites) was introduced to Lake of the Woods, Ontario, by the introduction of Smallmouth Bass (Armstrong 1985). Similarly, a bait bucket introduction of the Red Shiner into the Colorado River drainage was believed responsible for introducing the Asian tapeworm (Bothriocephalus opsarichthydis) to that system (Dextrase and Coscarelli 1999).

In Europe, native populations of crayfish have been decimated by a fungal plague carried by non-indigenous crayfish from North America (Alderman et al. 1990, Lodge et al. 2000a). Zebra Mussels were also responsible for introducing a fish helminth parasite, Phyllodistomum folium, into the Ebro River, Spain (Peribáñez 2009).

Drake (2011) concluded that the bait pathway had the potential to spread disease and pathogens to new waters. Based on a statewide risk analysis, Phelps et al. (2014) concluded that movement of baitfish represented a high risk of spreading Viral Hemmoraghic Septicemia (VHS). Many Great Lakes jurisdictions, including Ontario, implemented restrictions on the movement of live baitfish in an attempt to minimize the spread of VHS (Koenig et al. 2009). Sutherland et al. (2004) believed that the harvest and transfer of wild baitfish posed a serious threat of spreading Heterosporosis into inland waters in the Great Lakes basin.

HABITAT ALTERATION

Some species have the capability to modify habitat conditions in their new environment (Cucherousset and Olden 2011). For example, Common Carp can destroy aquatic vegetation and decrease water clarity (Fuller et al. 1999). Bain (1993) concluded that Grass Carp could significantly alter habitat and associated biological resources. Invasive crayfish have been found to reduce macrophyte growth and have a strong impact on macrophtye re-establishment (Bakker and Dorenbosch 2013). The range expansion of Rusty Crayfish has been responsible for a reduction in aquatic macrophytes in some Wisconsin lakes (Olsen et al. 1991) and northwestern Ontario (Momot 1992). Lodge and Lorman (1987) found that Rusty Crayfish had the ability to restructure the macrophyte and invertebrate communities in waters they inhabited.

Alteration of water quality has been attributed to some aquatic invasive species. In New Zealand, Rowe (2004) attributed the decline in water clarity to the presence of several invasive fishes including Rudd, Tench, Goldfish, Yellow Perch, Carp, and catfish. Once established, Zebra Mussels have been found to reduce cholorophyll a and nutrient concentrations in a waterbody (Foy et al. 2004) as well as an increase in water clarity (Aldridge et al. 2004).

Golden Shiners (Notemigonus crysoleucas). Photo: Andrew Drake


GENETIC IMPACTS

Many species, utilized by anglers as bait, readily hybridize in the wild (Table 2). Intraspecific and interspecific hybridization can lead to reduced fitness of the original population. Hybridization with other native species can lead to genetic introgression. Genetic effects resulting from the hybridization of native and non-native trout has been documented to be a threat to native salmonids in western North America (Dunham et al. 2004).

The aggressive Red Shiner was noted to dilute the gene pool of native Cyprinella by hybridization (Fuller et al. 1999). Massive hybridization of the introduced Arroyo Chub with native tui chub was believed to have replaced all of the pure stock of Tui Chub in the Mohave River drainage, California (Hubbs and Miller 1943). Where introduced, Longnose Dace are known to hybridize with native Speckled Dace (Fuller et al. 1999). Rudd are known to readily hybridize with native Golden Shiner (Fuller et al. 1999).

Table 2. Examples of hybridization among baitfish species (from Kerr and Grant 2000).

HYBRID REFERENCE(S)

Arroyo Chub x Tui Chub Hubbs and Miller (1943)

Banded Darter x Tesselated Darter Raesly and Stauffer (1990)

Bluntnose minnow x Bullhead minnow Trautman (1981)

Bluntnose Minnow x Fathead Minnow Trautman (1981)

Central Stoneroller x Rainbow Shiner Dahlberg (1968)

Cutiips Minnow x Tonguetied Minnow Fuller and Nico (1996)

Common Shiner x Central Stoneroller Menzel (1978)

Common Shiner x Crescent Shiner Menzel (1977)

Common Shiner x Longnose Dace Ross and Cavender (1977)

Common Shiner x Rosyface Shiner Raney (1940), Miller (1963)

Crescent Shiner x Mountain Redbelly Dace Hambrick (1977)

Fallfish x Common Shiner Scott and Crossman (1973), Fuller et al. (1999)

Golden Shiner x Rudd Burkhead and Williams (1991), Fuller et al. (1999)

Lake Chub x Longnose Dace Nelson (1973)

Longnose Dace x Speckled Dace Fuller et al. (1999)

Mimic Shiner x Common Shiner Moyle (1973)

Mountain Sucker x Bluehead Sucker Sigler and Miller (1963)

Northern Redbelly Dace x Finescale Dace New (1962), Joswiak et al. (1982), Joswiak et al. (1985), Das and Nelson (1989)

Orangethroat Darter x Logperch Hubbs and Strawn (1957)

Owens Sucker x Santa Ana Sucker Fuller et al. (1999)

Red Shiner x Spotfin Shiner Page and Smith (1970)

River Chub x Longnose Dace Stauffer et al. (1979)

Sheepshead Minnow x Commanche Spring Pupfish Stevenson and Buchanan (1973)

Sheepshead Minnow x Leons Springs Pupfish Echelle and Echelle (1997)

Sheepshead Minnow x Pecos Pupfish Echelle and Connor (1989), Wilde and Echelle (1997)

Southern Redbelly Dace x Common Shiner Greenfield et al. (1973)

Wakasagi x Delta Smelt Dill and Cordone (1997)

White Shiner x Rosefin Shiner Menzel (1978)

Yellowfin Shiner x Bandfin Shiner Menzel (1978)


COMMUNITY ALTERATION

The introduction and establishment of a non-indigenous species can disrupt the entire biotic community including species of fish, insects, amphibians and vegetation (see reviews by Lodge et al. 2000 and Dunham et al. 2004). The spread of invasive species can replace locally or regionally unique organisms. The result can often be the loss of biodiversity and a homogenization of species assemblages (Rahel 2000, Scott and Helfman 2001).

Introductions of non-indigenous fish into southern Nevada, USA, resulted in the decline or elimination of three resident fish species (Deacon et al. 1964). The introduction of Flathead Catfish to several southern U.S. states was implicated with the decline of several native bullheads and sunfishes (Guire et al. 1984). The disappearance of the native Rio Grande Silvery Minnow was attributed to displacement by the introduced Plains Minnow (Fuller et al. 1999). Kuda and Griffith (1993) described how native Mottled Sculpin were displaced to different habitats after the establishment of Shoshone Sculpin. In the western United States, the introduction of Black Bullhead resulted in predation upon endangered Humpback Chubs and newly hatched sport fish as well as being responsible for the decline of the Chiracahua Leopard Frog (Fuller et al. 1999). Marsden and Jude (1995) reported reductions in the populations of Mottled Sculpin and Slimy Sculpin in areas where Round Gobies had become established. The introduction and spread of the Bull Chub in the James River, Virginia, was linked to the decline and disappearance of the native River Chub (Fuller et al. 1999). In Colorado, the non-native White Sucker is replacing native suckers in the Rio Grande and Colorado River drainages (Walker 1993).

Habitat degradation and the introduction of non-indigenous species were identified as the two most significant threats to imperiled fishes in North America (Wilcover et al. 1998, Jelks et al. 2008). Flather et al. (1994) reported that alien invasive species affected more than 35% of listed imperiled species in the United States. Lovell and Stone (2005) reported that, in the United States, 44 native species were endangered and an additional 27 were negatively affected due to introductions of non-indigenous species. In Canada, Dextrase and Mandrak (2006) reported that aquatic invasive species affected 26 of 41 listed species at risk with an additional 27 being negatively affected. Establishment of non-native Fathead Minnows has contributed to the decline of imperiled species including the Lost River Sucker and Shortnose Sucker in Oregon (U. S. Fish and Wildlife Service 1993) as well as the Chiracahua Leopard Frog in Arizona (Rosen et al. 1995). Introduced channel catfish were believed to exert a major negative effect

on the populations of several endangered species in the United States (Fuller et al. 1999). Introduction of the Red River Shiner to the Cimarron River system, Kansas and Oklahoma, displaced the rare and native Arkansas River Shiner (Fuller et al. 1999).

ECONOMIC IMPACTS

The economic costs of freshwater aquatic invasive species in the United States has been estimated at $4.1 billion annually (Pimentel et al. 1999). MacIsaac (2003) estimated costs of up to $750 million annually for damage to aquatic ecosystems in Canada. Colautti et al. (2006) estimated the costs of ten non-indigenous species in Canadian fisheries, agriculture and forestry totalled $187 million per year. Based on a 2004 survey of power generation and water treatment facilities, Connelly et al. (2006) estimated that, between 1989-2004, Zebra Mussels had an estimated $268 million dollar impact in North America.

DISPLACEMENT OF INDIGENOUS SPECIES

Miller et al. (1989) reported that detrimental effects of introduced species were identified in 38% of extinctions of North American fishes.

The introduction of Mosquitofish was associated with the extinction of Amistad Gambusia and San Marcos Gambusia (Miller et al. 1989). Introductions of several non-indigenous species including Mosquitofish was identified as the reason for the decline of native Banff Longnose Dace (Goodchild 1999).

Non-indigenous crayfish introductions have resulted in the loss of many native crayfish populations in Europe (Lodge et al 2000b). The Rusty Crayfish has displaced two native crayfish species in Ohio and Illinois (Lodge et al. 2000) as well as Wisconsin (Olsen et al. 1991). Lodge et al. (2000) concluded that the greatest single threat to crayfish biodiversity was the introduction of non-indigenous crayfish.

Native unionids in Lake St. Clair disappeared shortly after the establishment of the invasive Zebra Mussel (Nalepa 1994). Similarly, the introduction of Zebra Mussels resulted in the decline and extirpation of native freshwater mussels in the Rideau River, Ontario (Martel et al. 2001). All five species of native snail fauna of El Cariño de la Montaña, Mexico, were lost due to the invasion by the non-indigenous Screw Snail (Contreras-Arquitea and Contreras-Balderas 1999).


ADVERSE EFFECTS ON COMMERCIAL AND RECREATIONAL FISHERIES

Invasive species transform fisheries with changes that are rarely unidirectional due to uneven ecological changes and corresponding uneven changes for stakeholders because of the segregated nature of resource use. Despite these uneven changes, clear losers (species and stakeholders) are common themes. The increased ecological uncertainty imposed by invasions often makes meeting long-term fishery management objectives difficult (Drake and Mandrak In Press).

Bait harvester emptying leech traps. Photo: Brenda Koenig

Although difficult to quantify, the introduction of aquatic invasive species can have deleterious impacts on native fish stocks and the recreational fisheries which they support. For example, the invasion of the upper Great Lakes by Sea llmprey had devastating effects on the lake trout fishery (Smith 1968, Coble et al. 1990, Lupi et al 2003). The Great Lakes commercial and recreational fisheries for both native and introduced species would be severely reduced without the continued and expensive control of Sea Lamprey. In Europe, the establishment of non-indigenous crayfish resulted in reduced economic viability of native crayfish fisheries (Lodge et al. 2000a). Harvest for human consumption is now only 5% of historic levels (Lodge et al. 2000b).


Acknowledgements

I thank Jason Borwick, Jeff Brinsmead, David Copplestone, Andrew Drake, Scott Gibson and Brenda Koenig for constructive comments on an earlier draft of this document.

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Personal Communications

Drake, Andrew. Great Lakes Laboratory for Fisheries and Aquatic Sciences. Department of Fisheries and Oceans. Burlington, Ontario.

Koenig, Brenda. Biodiversity Branch. Ontario Ministry of Natural Resources. Peterborough, Ontario.

Robbins, Mark. Enforcement Branch. Ontario Ministry of Natural Resources. Peterborough, Ontario.

Websites

Delivering Aquatic Invasive Species in Europe

http://www.europe-aliens.org/aboutDAISIE.do

Global Invasive Species database

http://www.issg.org/database

Illinois Natural History Survey

http://wwx.inhs.illinois.edu/resources/iisg/organisms-in-trade/pathways/livebait

United States Environmental Protection Agency

http://water.epa.gov/type/oceb/habitat/pathways.cfm

Additional References

GENERAL

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Boileau. M. G. 1985. The expansion of white perch (Morone americana) in the lower Great Lakes. Fisheries 10:6-9.

Bolton, T.F. and W.M. Graham. 2006. Jellyfish on the rocks: Bioinvasion threat of the international trade in aquarium live rock. Biological Invasions 8:651-653.

Bossenbroek, J. M., C. E. Kraft, and J. C. Nekola. 2001. Prediction of long-distance dispersal using gravity models: zebra mussel invasion of inland lakes. Ecological Applications 11:1778-1788.

Bothwell, M.L., D. R. Lynch, J. Deniseger, and H. Wright. 2009. Blooms of Didymosphenia geminata in rivers on Vancouver Island 1990 to present: A new invasive species. Journal of Phycology 45(Supplement 1):33-33.

Branstrator, D.K., M. E. Brown, L. J. Shannon, M. Thabes, and K. Heimgartner, K. 2006. Range expansion of Bythotrephes longimanus in North America: Evaluating habitat characteristics in the spread of an exotic zooplankter. Biological Invasions 8:1367–1379.

Brittan, M. R., J. D. Hopkirk, J. D. Connors, and M. Martin. 1970. Explosive spread of the oriental goby (Acanthogobius favimanus) in the San Franciso Bay-Delta region of California. Proceedings of the California Academy of Science 38:207-214.

Brown, B. 1993. Maine’s baitworm fisheries: Resources at risk? American Zoologist 33:568–577.

Brown, W. H. 1953. Introduced fish species in the Guadalupe River basin. Texas Journal of Science 5:245-251.

Burr, B. M. and R. J. Mayden. 1980. Dispersal of rainbow smelt (Osmerus mordax) into the upper Mississippi River. American Midland Naturalist 104:198-201.

Carlton, J. T. 1992. Introductions of marine and estuarine molluscs: An end of the 20th century perspective. Journal of Shellfish Research 11:489-505.

Carlton, J.T. 1996. Biological invasions and cryptogenic species. Ecology 77:1653–1655.

Carlton, J. T. 2001. Introduced species in the US coastal waters: Environmental impacts and management priorities. Pew Oceans Commission. Arlington, Virginia. 36 p.

Chang, A. L., J. D. Grossman, T. S. Spezio, H. W. Weiskiel, J. L. Blum, J. W. Burt, A. A. Muir, J. Piovia-Scott, K. E. Veblen and E. D. Grosholz. 2009. Tackling aquatic invasions: Risks and opportunities for the aquarium fish industry. Biological Invasions 11:773-785.

Coblentz, B. E. 1990. Exotic organisms: A dilemma for conservation biology. Conservation Biology 4:261-265.

Cohen, J., N. Mirotchnick and B. Leung. 2007. Thousands introduced annually: The aquarium pathway for non-indigenous plants to the St. Lawrence seaway. Frontiers in Ecology and the Environment 5:528-532.

http://www.europe-aliens.org/aboutDAISIE.do

http://www.issg.org/database/

http://wwx.inhs.illinois.edu/resources/iisg/organisms-in-trade/pathways/livebait/

http://wwx.inhs.illinois.edu/resources/iisg/organisms-in-trade/pathways/livebait/



Cohen, A. N., A. Weinstein, M. A. Emmett, W. Lau and J. T. Carlton. 2001. Investigations into the introduction of non-indigenous marine organisms via the cross-continental trade in marine baitworms. A report for the U.S. Fish and Wildlife Service, San Francisco Bay Program. 29 p.

Copp, G. H., C. Vaughn and A. Wheeler. 1993. First occurrence of the North American white sucker (Catostomus commersoni) in Great Britain. Journal of Fish Biology 42: 615-617.

Copp, G. H., L. Vilizzi and R. E. Gordon. 2010. The demography of introduction pathways, propagule pressure and occurrences of non-native freshwater fish in England. Aquatic Conservation Marine and Freshwater Ecosystems 20:595-601.

Courtenay, W. R. and J. E. Deacon. 1982. Status of introduced fishes in certain spring systems in southern Nevada. Great Basin Naturalist 42:361-366.

Courtenay, W. R. and J. E. Deacon. 1983. Fish introductions in the American southwest: A case history of Rogers Spring, Nevada. Southwestern Naturalist 28:221-224.

Courtenay, W. R. and J. R. Stauffer [eds.]. 1984. Distribution, biology and management of exotic fishes. Johns Hopkins. Baltimore, Maryland.

Crivelli, A.J. 1995. Are fish introductions a threat to endemic freshwater fishes in the northern Mediterranean region? Biological Conservation 72:311–319.

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Dalmazzone, S. 2000. Economic factors affecting vulnerability to biological invasions. p 17-30 In C. Perrings, M.H. Williamson, and S. Dalmazzone [eds.]. Economics of Biological Invasions, Edward Elgar Publishers, Cheltenham, England.

DeSilva, S. S., T. T. T. Nguyen, G. M. Turchini, U. S. Amarasinghe and N. W. Abery. 2009. Alien species in aquaculture and biodiversity: A paradox in food production. Ambio 38:24-28.

Duggan, I. C., C. A. M. Rixon, and H. J. MacIsaac. 2006. Popularity and propagule pressure: Determinants of introduction and establishment of aquarium fish. Biological Invasions 8:377-382.

Dundee, D. S. 1974. Catalog of introduced molluscs of eastern North America. Sterkiana 55:1–37.

Easton, R. S., D. J. Orth, and N. M. Burkhead. 1993. The first collection of rudd (Scardinius erythrophthajmus) in the New River, West Virginia. Journal of Freshwater Ecology 8:263-264.

Evans, M. S. 1988. Bythotrephes cederstsroemi: It’s new appearance in Lake Michigan. Journal of Great Lakes Research 14:234-240.

Franzin, W. G. 2003. Return to the sea: Homo sapiens help Osmerus mordax invade Hudson Bay. Paper presented at the 12th International Conference on Aquatic Invasive Species. July 9-12, 2003. Windsor, Ontario.

Gido. K. B and J. H. Brown. 1999. Invasion of North American drainages by alien fish species. Freshwater Biology 42:387-399.

Griffiths, R. W., D. W. Schloesser, J. H. Leach and W. P. Kovalak. 1991. Distribution and dispersal of the zebra mussel (Dreissena polymorpha) in the Great Lakes region. Canadian Journal of Fisheries and Aquatic Sciences 48:1381-1388.

Gunderson, J. L. and R. E. Kinnunen. Undated. The HACCP approach to prevent the spread of aquatic invasive species by aquaculture and baitfish operations. Sea Grant Publication. Duluth, Minnesota.

Hall, S. R. and E. L. Mills. 2000. Exotic species in large lakes of the world. Aquatic Ecosystems Health and Management 3:105-135.

Herbold, B. and P. B. Moyle. 1986. Introduced species and vacant niches. American Naturalist 128:751-760.

Hida, T. S. and D. A. Thomson. 1962. Introduction of the threadfin shad to Hawaii. Progressive Fish Culturist 24:159-163.

Holcik, J. 1991. Fish introductions in Europe with particular reference to its central and eastern part. Canadian Journal of Fisheries and Aquatic Sciences 48(Supplement 1): 945-957.

Howells, R. G. 1991. An introduction of hybrid crappie in a south Texas plan reservoir. Management Data Series 65. Texas Parks and Wildlife Department. Austin, Texas.

Imler, R. L. Striped bass introductions. Colorado Fisheries Research Review 8:19-20.

Jennings, M. R. and M. K. Saki. Establishment of red shiner (Notropis lutrensis) in the San Joaquin valley, California. California Fish and Game 76:46-57.

Jia, Y., Y. Chen, C. A. Gouodie, B. A. Simco, and Q. Liu. 2007. Potential invasion risk of the introduced American shad to aquatic ecosystems in China. Acta Zoologica Sinica 53:625-629.

Johnson, L.E., and J. T. Carlton. 1996. Post-establishment spread in large-scale invasions: Dispersal mechanisms of the zebra mussel Dreissena polymorpha. Ecology 77: 1686-1690,

Johnson, P. T. J., J. D. Olden, and M. J. Vander Zanden. 2008. Dam invaders: Impoundments facilitate biological invasions into freshwaters. Frontiers in Ecology and the Environment 6:357-363.

Kilian, J. V. and M. Ashton. 2007. Rusty crayfish invades Maryland. Caryfish News 29:5.

Kilroy, C. 2004. A new alien diatom (Didymosphenia geminata): Its biology, distribution, effects and potential risks for New Zealand fresh waters. Report CHC2004-128. National Institute of Water and Atmospheric Research. Christchurch, New Zealand

Leprieur, F., O. Beauchard, S. Blanchet, T. Oberdorff and S. Brosse. 2008. Fish invasions in the world’s river systems: When natural processes are blurred by human activities. Plos Biology 6:404–410.

Levine, J.M. and C.M. D’Antonio. 2003. Forecasting biological invasions with increasing international trade. Conservation Biology 17:322–26.

Linn, J. D. and T. C. Frantz.1965. Introduction of the opossum shrimp (Mysis relicta) into California and Nevada. California Fish and Game 51:48-51.

Lodge, D. M. 1993. Biological invasions: Lessons learned for ecology. Trends in Ecology and Evolution 8:133-137.


MacDonald, F. 2003. A case study of Canada’s response to aquatic invasive species introductions. Paper presented at the 12th International Conference on Aquatic Invasive Species. July 9-12, 2003. Windsor, Ontario.

MacIsaac, H. J., I. A. Grigorvich, J. A. Hoyle, N. D. Yan and V. E. Panov. 1999. Invasion of Lake Ontario by the Ponto-Caspian predatory cladoceran Cercopagis pengoi. Canadian Journal of Fisheries and Aquatic Sciences 56:1-5.

Marshall, T. L. and R. P. Johnson. 1971. History and results of fish introductions in Saskatchewan, 1900-1969. Fisheries Report No. 8. Fish and Wildlife Branch. Saskatchewan Department of Natural Resources. Regina, Saskatchewan. 30 p.

Maxwell, W.D. 1974. A history of the California live-baitfishing industry. Marine Resources Technical Report No. 2. California Department of Fish and Game. Long Beach, California.

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Meyerson, L. A. and H. A. Mooney. 2007. Invasive alien species in an era of globalization. Frontiers in Ecology and the Environment 5:199–208.

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Moyle, P. B. 1986. Fish introductions into North America: Patterns and ecological impact. p. 27-43 In H. A. Mooney and J. A. Drake [eds.]. Ecology of Biological Invasions of North America and Hawaii. Springer- Verlag Press. New York, New York.

Moyle, P. B. and T. Light. 1996. Biological invasions of fresh water: Empirical rules and assembly theory. Biological Conservation 56:149-161.

Moyle, P.B. and T. Light. 1996. Fish invasions in California: Do abiotic factors determine success? Ecology 77:1666–1670.

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Nielsen, L. A. 1982. The baitfish industry in Ohio and West Virginia with special reference to the Ohio River sport fishery. North American Journal of Fisheries Management 2:232-238.

Pelzman, R. J. 1972. Evaluation of the introduction of Tilapia sparrmanii into California. Inland Fisheries Report 72-3. California Department of Fish and Game. Sacramento, California.

Peterson, D. L. and F. A. Hennagir. 1980. Minnesota live bait industry assessment study. Completion Report. Commercial Fisheries Research and Development Project. National Marine Fisheries Service. Minneapolis, Minnesota.

Pierce, J. M. and M. E. Wachtman. 1971. The live bait industry in Ohio: It’s extent, condition and problems. Completion Report. Commercial Fisheries Research and Development Project. National Marine Fisheries Service. Columbus, Ohio.

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Radar, R. B., M. C. Belk and M. J. Keleher. 2003. The introduction of an invasive snail (Melanoides tuberculata) to spring ecosystems of the Bonneville basin, Utah. Journal of Freshwater Ecology 18:647.

Ricciardi, A. and H. J. MacIsaac. 2000. Recent mass invasion of the North American Great Lakes by Ponto-Caspian species. Trends Ecological Evolution 15:62-65.

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Ruesink, J. L. 2005. Global analysis of factors affecting the outcome of freshwater fish introductions. Conservation Biology 19:1883-1893.

Shaw, B. and A. Howell. 2013. Bait shop “toolkit” and outreach: Engaging bait shops as opinion leaders in the prevention of aquatic invasive species. Paper presented at the 18th International Conference on Aquatic Invasive Species. April 21-25, 2013. Niagara Falls, Ontario.

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Strayer, D. L. 2009. Twenty years of zebra mussels: Lessons from the mollusk that made headlines. Frontiers in Ecology and the Environment 7: 135–141

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Vitousek, P. M. 1990. Biological invasions and ecosystem process: Towards an integration of population biology and ecosystem studies. Oikos 57:7-13.

Welcomme, R. L. 1991. International introductions of freshwater fish species into Europe. Finnish Fisheries Research 12:11-18.

Werner, G. 1972. Bluespotted sunfish (Enneacanthus gloriosus) in the Lake Ontario drainage, New York. Copeia 1972:878-879.

Westphal, M. I., M. Browne, K. MacKinnon and I. Noble, I. 2008. The link between international trade and the global distribution of invasive alien species. Biological Invasions 10:391–398.

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VECTORS AND PATHWAYS OF AQUATIC INVASIVE SPECIES

Balcom, N. and C. Yarish. 2009. Don’t dump bait: Marine bait worms as a potential vector of non-native species. Connecticut Sea Grant Publication. Groton, Connecticut. 2 p. http://web2.uconn.edu/seagrant/whatwedo/ais/btwrms.pdf

Brock, V. E. 1960. The introduction of aquatic animals into Hawaiin waters. Internationale Revue der Gesamten Hydrobiolgie 45:463-480.

Carlton, J. T. 1993. Dispersal mechanisms of the zebra mussel (Dreissena polymorpha). p. 677-697 In T. F. Nalepa and D. W. Schloesser [eds.]. Zebra Mussels: Biology, Impacts and Control. Lewis Publishers. Boca Raton, Florida.

Courtenay, W. R. 1993. Biological pollution through fish introductions. p. 35-61 In B. N. McKnight [ed.]. Biological Pollution: The Control and Impact of Invasive Aquatic Species. Indiana Academy of Science. Indianapolis, Indiana.

Cowie, R. H. and D. G. Robinson. 2003. Pathways of introduction of freshwater snails and slugs. p. 93- 122 In G. M. Ruiz and J. T. Carlton [eds.]. Invasive Species: Vectors and Management Strategies. Island Press. Washington, D. C.

DiStefano, R. J., M. E. Litvan and P. T. Horner. 2008. The bait industry as a potential vector for alien crayfish introductions: Problem recognition by fisheries agencies and a Missouri evaluation. Missouri Department of Conservation Final Report, Columbia. Missouri.

Dzialowski, A. R., W. J. O’Brien W. M. Swaffer. 2000. Range expansion and potential dispersal mechanisms of the exotic cladoceran Daphnia lumholtzi. Journal of Plankton Research 22:2205-2223.

Floerl O. and G. J. Inglis. 2005. Starting the invasion pathway: The interaction between source populations and human transport vectors. Biological Invasions 7: 589-606,

Gertzen, E., O. Familiar and B. Leung. 2008. Quantifying invasion pathways: Fish introductions from the aquarium trade. Canadian Journal of Fisheries and Aquatic Sciences 65:1265-1273.

Heard, W. R. 1956. Live bait imports: Chrosomus eos and Eucalia inconstans as potential additions to Oklahoma’s fish fauna. Proceedings of the Oklahoma Academy of Science 1956:47-48.

Henttonen P, and J. Huner. 1999. The Introduction of alien species of crayfish in Europe: A historical introduction. Crustacean Issues 11:13-22.

Hobbs, H. H., J. P. Jass and J. V. Huner. 1989. A review of global crayfish introductions with particular emphasis on two North American species (Decapoda, Cambaridae). Crustaceana 56:299–316

Hulme, P. E., S. Bacher, M. Kenis, S. Klotz, I. Kuhn, D. Minchin, W. Nentwig, S. Olenin, V. Panov, J. Pergl, P. Pybek, A. Roques, D. Sol, W. Solarz and M. Vila. 2008. Grasping at the routes of biological invasions: A framework for integrating pathways into policy. Journal of Applied Ecology 45:403–414.

Johnson, L. E. and J. M. Bossenbroek and C. E. Kraft. 2006. Patterns and pathways in the post-establishment spread of non-indigenous aquatic species: The slowing invasion of North American inland lakes by the zebra mussel. Biological Invasions 8:475-489.

Kilian, J. V., R. J. Klauda, S. Widman, M. Kashiwagi, R. Weglein and S. Schuster. 2010. A study of the sale, use and release of live bait in Maryland: A call for stricter regulations, better education, and enhanced monitoring to reduce the threat of invasive species introductions. Report prepared for the Maryland Department of Natural Resources. Annapolis, Maryland.

Lau, W. 1995. Importation of baitworms and shipping seaweed: Vectors for introduced species? p 21–38 In D. Sloan, M. Christensen, and D. Kelso [eds.]. Environmental Issues: From a Local to a Global Perspective. Environmental Sciences Group Major. University of California. Berkeley, California.

LoVullo, T. J., and J. R. Stauffer, Jr. 1993. The retail bait-fish industry in Pennsylvania – source of introduced species. Journal of the Pennsylvania Academy of Science 67(1):13–15.

Ludwig, H. R. 1995. Bait bucket transfer potential between the Mississippi and the Hudson Bay watersheds. M.Sc. Thesis. North Dakota State University. Fargo, North Dakota.

McDowell, R. M. 2004. Shoot first and then ask questions: A look at aquarium imports and invasiveness in New Zealand. New Zealand Journal of Marine and Freshwater Research 38:503-510.

Mullady, T.L., T.K. Rawlings and G.M. Ruiz. 2000. Importation of organisms associated with baitworms from Vietnam. p. 402 In J. Pederson, J. [ed.]. Proceedings of a Marine Bioinvasions Conference. January 24-27,1999. MIT Sea Grant College Program, Massachusetts Institute of Technology, Cambridge, Massaachusetts.

Olson, A. M. 2001. Do live marine products serve as pathways for the introduction of non-indigenous species? p. 243–247 In B. C. Paust and A. A. Rice [eds.]. Marketing and Shipping Live Aquatic Products: Proceedings of the Second International Conference and Exhibition. November, 1999. Seattle, Washington.

Passarelli, B. 2010. The marine live bait trade in California: A pathway for introduction of non-indigenous species? M.Sc. Thesis. California State University. Long Beach, California.

Picco, A. M. and J. P. Collins. 2008. Amphibian commerce as a likely source of pathogen pollution. Conservation Biology 22:1582-1589.

Puth, L. M. and T. F. H. Allen. 2005. Potential corridors for the rusty crayfish, Orconectes rusticus, in northern Wisconsin, USA, lakes: Lessons for exotic invasions. Landscape Ecology 20:567–577.

Ruiz, G. M., and J. T. Carlton. 2003. Invasion vectors: A conceptual framework for management. In G. M. Ruiz and J. T. Carlton [eds.]. Invasive species: Vectors and Management Strategies. Island Press, Washington, D.C.

Secretariat of the Convention on Biological Diversity. 2010. Pets, aquarium and terrarium species: Best practices for addressing risks to biodiversity. Montreal, Québec.

Sherfy, M. 2000. International trade in live bait: A potential source of aquatic nuisance species introduction. In Proceedings of the Non-indigenous Species of the Mid-Atlantic Workshop, June 2000.

Weigle, S.M., L. D. Smith, J. T. Carlton, and J. Pederson. 2005. Assessing the risk of introducing exotic species via the live marine species trade. Conservation Biology 19:213–223

http://web2.uconn.edu/seagrant/whatwedo/ais/btwrms.pdf

http://web2.uconn.edu/seagrant/whatwedo/ais/btwrms.pdf


Wilson, J. R. U., E. E. Dormontt, P. J. Prentis, A. J. Lowe, and D. M. Richardson. 2009. Something in the way you move: Dispersal pathways affect invasion success. Trends in Ecology and Evolution 24:136-144.

ECOLOGICAL IMPACTS OF INTRODUCTIONS OF NON-INDIGENOUS SPECIES

Achieng, A. P. 1990. The impact of the introduction of Nile perch (Lates niloticus) on the fisheries of Lake Victoria. Journal of Fish Biology 37:17-23.

Adams, C. E. and P. S. Maitland. 1998. The ruffe population of Loch Lomond, Scotland: It`s introduction, population expansion and interaction with native species. Journal of Great Lakes Research 24:249-262.

Amundsen, P. A., A. Siwertsson, R. Primicercio and T. Bohn. 2009. Long term responses of zooplankton to invasion by a planktivorous fish in a subarctic watercourse. Freshwater Biology 54:24-34.

Anastacio, P. M. and J. C. Marques. 1997. Crayfish (Procambarus clarkii) effects on initial stages of rice growth in the lower Mondego River valley. Freshwater Crayfish 11:608-617.

Axelsson, E., P. Nystrom, J. Sidenmark and C. Bronmark. 1997. Crayfish predation on amphibian eggs and larvae. Amphib-Reptilia 18:217-228.

Ayala, J. R., R. B. Rader, M. C. Belk and G. B. Schaalje. 2007. Ground truthing the impact of invasive species: Spatio-temporal overlap between native least chub and introduced western mosquitofish. Biological Invasions. 9:857-869.

Bain, M. B., E. R. Cornwell, K. M. Hope, G. E. Eckerlin, R. N. Casey, G. H. Groocock, R. G. Getchell, P. R. Bowser, J. R. Winton, W. N. Batts, A. Cangelosi and J. W. Casey. 2010. Distribution of an invasive aquatic pathogen (Viral Hemorrhagic Septicemia Virus) in the Great Lakes and its relationship to shipping. PLOS One 5:1-8.

Bartley, D.M. and C.V. Casal. 1999. Impact of introductions on the conservation and sustainable use of aquatic biodiversity. FAO Aquaculture Newsletter 20: 15-19.

Bauman, J. R. and T. J. Kwak. 2011. Trophic relations of introduced flathead catfish in an Atlantic river. Transactions of the American Fisheries Society 140:1120-1134.

Baxter, C. V., K. D. Fausch, M. Murakami and P. L. Chapman. 2007. Invading rainbow trout usurp a terrestrial prey subsidy from native charr and reduce their growth and abundance. Oecologia 153:461-470.

Beers, J. H. and R. McConnell. 1966. Some effects of threadfin shad introduction on black crappie diet and condition. Journal of the Arizona Academy of Science 4:71-74.

Bergman, E. 1990. Effects of roach (Rutilis rutilis) on two percids (Perca fluviatilis and Gymnocephaous cernua): Importance of species interactions for diet shifts. Oikos 57:241-249.

Biro, P. 1977. Effects of exploitation, introductions and eutrophication on percids in Lake Balaton. Journal of the Fisheries Research Board of Canada 345:1678-1683.

Blanco, S. and L. Ector. 2009. Distribution, ecology, and nuisance effects of the freshwater invasive diatom Didymosphenia geminata: A literature review. Nova Hegwigia 88:347-422.

Bradford, D. F. 1989. Allopatric distribution of native frogs and introduced fishes in high Sierra Nevada lakes of California: Implications of the negative effect of fish introductions. Copiea 1989:775-778

Bradford, D. F., T. Tabatabai and D. M. Graber. 1993. Isolation of remaining populations of the native frog Rana muscosa by introduced fishes in Sequoia and Kings Canyon National Parts, California. Conservation Biology 7:882-888.

Bradford, D. F., S. D. Cooper, T. M. Jenkins, K. Kratz, O. Sarnelle and A. D. Brown. 1998. Influences of natural acidity and introduced fish on faunal assemblages in California alpine lakes. Canadian Journal of Fisheries and Aquatic Sciences 55:2478-2491.

Branstrator, D. K. 1995. Ecological interactions between Bythotrephes cederstroemi and Leptodora kindti and the implications for species replacement in Lake Michigan. Journal of Great Lakes Research 21:670-679.

Brey, M. K. 2012. Trophic interactions of introduced fishes in a reservoir food web. Ph.D. Dissertation. North Carolina State University. Raleigh, North Carolina.

Britton, J. R., J. Cucherousset, J. Grey and R. E. Gozlan. 2011. Determining the strength of exploitive competition from an introduced fish: Role of density, biomass and body size. Ecology of Freshwater Fish 20:74-79.

Brown, L. R. and P. B. Moyle. 1981. The impact of squawfish on salmonid populations: A review. North American Journal of Fisheries Management 1:104-11.

Bunnell, D. B., T. B. Johnson and C. T. Knight. 2005. The impact of introduced round gobies (Neogobius melanostomus) on phosphorus cycling in central Lake Erie. Canadian Journal of Fisheries and Aquatic Sciences 62:15-29.

Burkhardt, S. and J. T. Lehman. 1994. Prey consumption and predatory effects of an invertebrate predator (Bythotrephes) based on a phosphorus budget. Limnology and Oceanography 39:1007-1019.

Byers, J. E. and L. Goldwasser. 2001. Exposing the mechanism and timing of impact of non-indigenous species on native species. Ecology 82:1330-1343.

Cadwallader, P. L. 1996. Overview of the impacts of introduced salmonids on Australian native fauna. Australian Nature Conservation Agency. Canberra, Australia.

Cahn, A. J. 1983. The effect of carp on a small lake: The carp as dominant. Ecology 10:371-374.

Cambray, J. A. 2003. Impact on indigenous species biodiversity caused by globalization of alien recreational freshwater fisheries. Hydrobiologia 500:217-230.

Capelli, G. M. 1982. Displacement of northern crayfish by Orconectes rusticus. Limnology and Oceanography 27:741-745.

Capelli, G. M. and B. L. Munjal. 1982. Aggressive interactions and resource competition in relation to species displacement among crayfish of the genus Orconectes. Journal of Crustacean Biology 2:486-492.

Carlson, R. L. 2008. Morphological change in the tessellated darter (Etheostoma olmstedi) follow the introduction of the banded darter (E. zonale) to the Susquehanna River drainage. Copeia 2008:661-668.


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Alien Species – species of plants, animals, and micro-organisms introduced by human action outside their natural past or present distribution.

Aquatic Species – an organism which lives in a waterbody or a wetland during all or part of their lives (from Benson 1999).

Bait Bucket Transfer – the transport and subsequent release of water and aquatic biota into a basin where it was previously absent.

Ecological Filters Concept – A concept which describes the successive arrival, survival, establishment and spread in determining invasion succesas or failure.

Establishment – the process of a non-native species in a new habitat successfully producing viable offspring with the likelihood of continued survival.

Human Assisted Dispersal – any anthropogenic mechanism, whether intentional or unintentional, which can result in dispersal of a species beyond its native range.

Invasive species – harmful alien organisms whose introduction or spread threatens the environment, the economy, or society.

Pathway – the ways in which invasive alien species are introduced or spread.

Vector – the means by which species from a source population follows a pathway to a new destination

Glossary


APPENDIX 1. Some of the more common potential pathways for the transfer and movement of aquatic invasive species in Ontario.

PATHWAY REFERENCE

Aquaculture escapes Mills et al. (1974), Welcomme (1988), Courtney and Williams (1992), Benson (1999), Crossman and Cudmore (1999a), Naylor et al. (2001), Kolar and Lodge (2002), Elvira and Almodovar (2005), Kerr et al. (2005), Molnar et al. (2008), Peters and Lodge (2009), Grosholz et al (2012), Kinnunen and Gunderson (2013), DFO (undated)

Aquarium and ornamental water industry Werner (1972), Welcomme (1988), Mills et al. (1994), Benson (1999), Courtenay and Stauffer (1990), Courtney (1999), Crossman and Cudmore (1999), Mackie (1999a), Carlton (2001), Kolar and Lodge (2002), Noel (2003), Lintermans (2004), Padilla and Williams (2004), Kerr et al. (2005), Rixon et al. (2005), Jensen et al. (2006), Keller and Lodge (2007), Gertzen et al. (2008), Chavanich et al. (2009), Peters and Lodge (2009), Duggan (2010), Strecker et al. (2011), Williams et al. (2012), Chucholl (2013), Coetzee and Martin (2013), Gettys et al. (2013),


Ballast water Welcomme (1988), Griffiths et al. (1991), Hulme (1993), Mills et al. (1993), Mills et al. (1994), Eno et al. (1997), Crossman and Cudmore (1999c), Mackie (1999b), Wiley and Claudi (1999), Carlton (2001), Kolar and Lodge (2002), Gollasch (2004), Holeck et al. (2004), Lintermans (2004), Mills et al. (2004), Niimi (2004), Kerr et al. (2005), Ricciardi (2006), Molnar et al. (2008), Bailey et al. (2009), Briski et al. (2009), Kipp et al. (2009), MacIsaac (2013), DFO (undated),


Recreational boating and fishing gear Johnstone et al. (1985),Padilla et al. (1996), Buchan and Padilla (2001), Johnson et al. (2001), Kerr et al. (2005), Leung et al. (2006), Bobeldyk et al. (2007), Jacobs and MacIsaac (2007), Gates et al. (2008), Vander Zanden and Olden (2008), Acosta and Forrest (2009), Bothwell et al. (2009), Darbyson et al. (2009), Timar and Phaneuf (2009), Rothlishberger et al. (2010), Ashton et al. (2012), Anderson et al. (2013), MacIsaac (2013), Sheehan and Lucy (2013), DFO (undated),


Cross basin connections Scott and Christie (1963), Emery (1985), Mills et al. (1993), Mills et al. (1994), Buchan and Padilla. (1999), Crossman and Cudmore (1999b), Dextrase and Coscarelli (1999), Fuller et al. (1999), Mills et al. (1999), Minns and Cooley (1999), Wright and Franzin (1999), Freeman et al. (2002), Kerr et al. (2005), Galil et al. (2007), Mandrak and Kim (2013), Modley (2013), DFO (undated),


Intentional introductions (legal and illegal) Holcomb (1964), Christie (1972), Moyle (1976), Emery (1985), Evans and Loftus (1987), Welcomme (1988), Crossman (1991), Mills et al. (1993), Mills et al. (1994), Dextrase and Coscarell (1999), Crawford (2001), Commission for Environmental Cooperation (2003), Rahel (2004), Elvira and Almodovar (2005), Kerr et al. (2005), Koel et al. (2005), Warner (2005), Boucher (2007), Johnson et al. (2009), Lapointe (2010), Sheehan and Lucy (2013), DFO (undated),








PATHWAY REFERENCE

Live food fish industry Welcomme (1988), Mackie (1999c), Kerr et al. (2005), Rixon et al. (2005), Cudmore and Mandrak (2006), Keller and Lodge (2007), Choudhury et al. (2013), MacIsaac (2013), DFO (undated),


Science/laboratory escapes Emery (1985), http://water.epa.gov/type/oceb/habitat/pathways.cfm

Use of bait by recreational anglers Welcomme (1988), Nearly and Leach (1992), Mills et al. (1994), Ludwig (1995), Ludwig and Leitch (1996), Gido and Brown (1999), Goodchild (1999), Litvak and Mandrak (1999), Hackman (2002), Kolar and Lodge (2002), Wallace (2003), Kerr et al. (2005), Bobeldyk et al. (2007), Keller and Lodge (2007), DiStefano et al. (2009), Peters and Lodge (2009), Drake and Mandrak (2010), Lapointe (2010), Feiner et al. (2013), Kilian and Klauda (2013), Lieb (2013), Drake and Mandrak (2014a, 2014b), DFO (undated),


Religious/cultural beliefs Lintermans (2004), Pathak (2013)

Waterfowl Yan et al. (1992), Shurin and Havel (2002), Sheehan and Lucy (2013)





APPENDIX 2. Introduction or spread of aquatic invasive species attributed to the recreational use of bait.

SPECIES WATERBODY YEAR(S) REFERENCE

Asian freshwater clam (Corbicula fluminea)

North America – Carlton (1992), Goodchild (1999)

Arroyo chub (Gila orcutti) California, USA 1930s Fuller et al. (1999)

American (rainbow) smelt (Osmerus mordax)

Ontario inland waters

Quetico Provincial Park, Ontario

– Litvak and Mandrak (1993), Mercado-Silva et al. (2006)

Goodchild (1999)

Arkansas River shiner (Notropis girardi) Pecos River (New Mexico) Washita River (Oklahoma)

1978

1960s

Bestgen et al. (1989)

Cross (1970)

Bandfin shiner (Luxilus zonistius) Alabama and Georgia, USA 1950s Fuller et al. (1999)

Bigeye chub (Hybopsis amblops) Lower Hudson River, Maine, USA 1987 Fuller et al. (1999)

Bigeye shiner (Notropis boops) Lower Missouri River, USA 1950s Pflieger (1997)

Blackbelly salamander (Desmognathus quadromaculatus)

Georgia, USA Benson (1999)

Blacknose dace (Rhinichthys atratulus) North Carolina, USA Fuller et al. (1999)

Black jumprock (Scartomyzon cervinus) Virginia, USA 1951 Fuller et al. (1999)

Blackside darter (Percina maculata) Wisconsin, USA – Fuller et al. (1999)

Bluehead chub (Nocomis leptocephalus) Alabama, Florida, North Carolina, and Virginia, USA

1940s Fuller et al. (1999)

Bluntnose minnow (Pimephales notatus) Connecticutt, Georgia, Massachusetts, Ohio, South Dakota, Utah, Virginia, West Virginia, USA

Lake St. Marys, Ohio

– Fuller et al. (1999)

Hubbs and Black (1947), Fuller et al. (1999)

Brassy minnow (Hybognathus hankinsoni) Colorado, New York, Tennessee and Utah, USA


Brook stickleback (Culaea inconstans) Several U.S. states – Fuller et al. (1999)

Brown darter (Etheostoma edwini) Florida, USA 1960s Fuller et al. (1999)

Bull chub (Nocomis raneyi) North Caroline and Virginia, USA – Fuller et al. (1999)

Bullhead minnow (Pimephales vigilax) Mexico – Contreras-Balderas (1999)

Calanoid copepod (Skistodiaptomus pallidus)

Lakes Erie, Ontario and St. Clair – Mills et al. (1993), Benson (1999)

California roach (Hesperoleucas symmetricus)

Oregon, USA –

Calico crayfish (Orconectes immunis) Central Europe

Several U.S. states

– Benson (1999), Chucholl (2013)

Benson (1999)

Central stoneroller (Campostoma anomalum)

Northeastern USA 1980s-90s Fuller et al. (1991)

Chub shiner (Notropis potteri) Red River, Oklahoma and Texas – Hall (1956)

Crayfish (Orconectes causeyi) New Mexico – Benson (1999)

Creek chub (Semotilus atromaculatus) Colorado, Texas, Utah and Wyoming, USA


Crescent shiner (Luxilus creasinus) North Carolina and Virginia, USA 1965



Cutlips minnow (Exoglossum maxillingua) Connecticutt, Virginia and West Virginia, USA


Common carp (Cyprinus carpio) Australia – Lintermans (2004)

Common shiner (Luxilis cornutus) West Virginia, USA – Fuller et al. (1999)

Cutlip minnow (Exoglossum maxillingua) New River drainage, Virginia and West Virginia, USA

– Goodchild (1999)

Eastern silvery minnow (Hybognathus regius)

Maine, USA 1982 Fuller et al. (1999)

Emerald shiner (Notropis atherinoides) New York and Maine, USA – Fowler (2013)

European green crab (Carcinus maenas) San Francisco bay, USA – Fowler (2013)

Fallfish (Semotilus corporalis) New York and Virginia, USA – Fuller et al. (1999)

Fathead minnow (Pimephales promelas) Thirty-five U.S. states

Mexico

– Benson (1999)

Contreras-Balderas (1999)

Fieryback shiner (Cyprinella pyrrhomelas) South Carolina, USA – Johnston et al. (1995), Fuller et al. (1999)

Flathead chub (Plagopterus argentissius) Arizona, USA 1950s Fuller et al. (1999)

Florida sand darter (Ammocrypta bifascia) Florida, USA 1970s Fuller et al. (1999)

Gammarid amphipod (Gammarus fasciatus)

Great Lakes basin – Mills et al. (1993)

Ghost shiner (Notropis buchanani) Great Lakes

(Ontario waters)

Ohio

1979 Mills et al. (1993), (1994)

Fuller et al. (1999)

Golden Shiner (Noptemigonus crysoleucas)

North America (including the Great Lakes)

Mexico

Two Prince Edward Island lakes

–

–

Benson (1999), Keller and Lodge (2007)


Goodchild (1999)

Goldfish (Carassius auratus) Australia

All U.S. states except Alaska

Great Lakes basin

Hungary, Europe

–

–

–

1891, 1954

Lintermans (2004)

Benson (1999)

Mills et al. (1993)

Welcomme (1988)

Green sunfish (Lepomis cyanellus) Northern tributary of Lake Ontario, Ontario

– Litvak and Mandrak (1999)

Gulf killifish (Fundulus grandis) New Mexico and Texas, USA – Fuller et al. (1999)

Highscale shiner (Notropis hypsilepsis) Savannah River drainage, Georgia – Fuller et al. (1999)

Hitch (Lavinia exilicauda) Southern California 1954 Fuller et al. (1999)

Leatherside chub (Gila copei) California, USA – Fuller et al. (1999)

Lined topminnow (Fundulus lineolatus) Florida 1950s Fuller et al. (1999)

Longfin dace (Rhinichthys chrysogaster) Arizona and New Mexico, USA – Fuller et al. (1999)

Longjaw mudsucker (Gillichthys mirabilis) Arizona, USA – Fuller et al. (1999)

Longnose dace (Rhinichthys cataractae) Colorado, Kentucky, Virginia and Wyoming, USA


Longnose shiner (Notropis longirastris) Tussahaw Creek, Georgia – Fuller et al. (1999)

Longnose sucker (Catostomus catostomus)

Colorado, Connecticut and Wyoming, USA




Margined madtom (Noturus insignis) Western Québec, Canada

Several northeastern U.S. states

Great Lakes basin

1971

–

Rubec and Coad (1974), Litvak and Mandrak (1993)


Mills et al. (1993)

Mexican tetra (Astyanax mexicanus) Arizona, California, Louisiana, New Mexico, Ohio and Oklahoma, USA

Mimic shiner (Notropis volucellus) Connecticutt, Massachusetts, Pennsylvania, West Virginia, USA

– Hartel (1992), Fuller et al. (1999)

Mosquitofish (Gambusia affinis) North America

Alberta

–

1924

Keller and Lodge (2007)

Goodchild (1999)

Mottled sculpin (Cottus bairdii) Arizona, California, Colorado, and Nevada, USA


Mountain sucker (Catostomus platyrhlynchus)

Arizona. Nevada, Utah, USA 1940s Fuller et al. (1999)

Mountain treefrog (Hyla eximia) Maryland and Texas, USA – Benson (1999)

Mozambique tilapia (Oreochromis mossambicus)

Australia – Lintermans (2004)

Mummichog (Fundulus heteroclitus) New Hampshire and Pennsylvania, USA


Ninespine stickleback (Pungitius pungitiius)

Colorado, USA – Walker (1993), Fuller et al. (1999).

Northern hog sucker (Hypentelium nigricans)

Alabama, Massachusetts and Tennesse, USA


Northern pearl dace (Margariscus margarita)

Connecticut, USA – Fowler (2013)

Northern studfish (Fundulus catenatus) Kentucky, Missouri, Oklahoma and West Virginia, USA


Orangefin madtom (Noturus gilberti) Virginia 1951 Fuller et al. (1999)

Oriental weatherfish (Misgurnus anguilliacaudatus)

Australia – Lintermans (2004)

Ozark minnow (Notropis nubilus) Oklahoma, USA – Miller and Robison (1973)

Ozark shiner (Notropis ozarcanus) Illinois River, Arkansas – Burr et al. (1979)

Periwinkel snail (Littorina saxatilis) San Francisco bay, USA – Fowler (2013)

Pirate perch (Aphredoderus sayanus) Wisconsin, USA – Fuller et al. (1999)

Plains killifish (Fundulus zebrinus) Mexico

Arizona, Colorado, Montana, New Mexico, South Dakota, Texas, Utah, and Wyoming, USA

– Contreras-Balderas (1999)


Plains minnow (Hybognathus placitus) Pecos River, New Mexico, USA 1968 Fuller et al. (1999)

Plains topminnow (Fundulus sciadicus) Colorado and Nebraska, USA – Fuller et al. (1999)

Popeye shiner (Notropis ariommus) West Virginia, USA – Fowler (2013)

Rainbow shiner (Notropis chrosomus) Black Warrior River and Town Creek, Alabama

South Chickamauga Creek, Georgia

–

–

Starnes and Etnier (1986)

Starnes and Etnier (1986)



Redeye chub (Notropis harperi) Ochlockonee River drainage, Florida


Redfin perch (Perca fluviatilis) Australia – Lintermans (2004)

Red shiner (Cyrpinella lutrensis) Alabama, Arizona, Colorado, Georgia, and Illinois, USA

Mexico



Red River shiner (Notropis bairdi) Kansas and Oklahoma, USA – Fowler (2013)

Redlip shiner (Notropis chiliticus) Little River system, North Carolina

Big Chestnut Creek, Virginia

Deep River system, North Carolina

1963

1976

1985

Richards and Carnes (1964)


Jenkins and Burkhead (1994)

Redside dace (Clinostomus elongatus) Maryland, USA 1970s Fuller et al. (1999)

Redside shiner (Richardsonius balteatus)

Arizona, Colorado, Montana, Utah and Wyoming, USA

Lees Lake, Alberta


Goodchild (1999)

Rio Grande sucker (Catostomus plebeius) Arizona, New Mexico, USA – Minckley (1973)

River chub (Nocomis micropogon) Eastern Ontario waters

Georgia, North Carolina, and South Carolina USA

– Goodchild and Tilt (1976)


River shiner (Notropis blennius) Bricken Spring, Oklahoma 1940s Hubbs and Bonham (1951)

Rosefin shiner (Lythrurus ardens) Alabama, North Carolina, Ohio, Virginia, USA

– Fowler (2013)

Rosyface shiner (Notropis rubellus) Connecticutt River, Connecticutt

Delaware River, Pennsylvania

–

–



Rough shiner (Notropis baileyi) Alabama and Florida, USA – Fowler (2013)

Rudd (Scardinius erythrophthalmus) Great Lakes drainage 1989 Klindt (1990), Crossman (1991), Crossman et al. (1992), Mills et al. (1993), Mills et al. (1994), Benson (1999), Fuller et al. (1999), Kapuscinski et al. (2012)

Rusty crayfish (Orconectes rusticus) Ontario and northern United States

Twenty-one U.S. states

– Benson (1999), Kerr et al. (2005), Keller and Lodge (2007), Berube and Kraft (2010), Hamr (2010),

Benson (1999)

Seminole killifish (Fundulus seminolis) Florida, USA 1960s Fuller et al. (1999)

Sharpnose shiner (Notropis oxyrhynchus) Colorado River, Texas 1950s Yurgens (1954)

Sheepshead minnow (Cyprinodon variegatus)

New Mexico and Texas, USA – Fuller et al. (1999)

Signal crayfish (Pacifastacus leniusculus) California, Nevada and Utah, Unites States

– Benson (1999)

Silverband shiner (Notropis shumardi) Oklahoma and Texas, USA 1950s Fuller et al. (1999)

Silverjaw minnow (Notropis buccatus) Ocmulgee River system, Georgia

Clinch River, Virginia

1970s

–



Slender madtom (Noturus exilis) Arkansas, USA – Robison and Winters (1978)

Slim minnow (Pimephales tenellus) Kansas, USA – Fuller et al. (1999)



Smalleye shiner (Notropis buccula) Colorado River, Texas – Fuller et al. (1999)

Southern studfish (Fundulus stellifer) Georgia, USA – Fuller et al. (1999)

Speckled dace (Rhinichthys osculus) California, USA – Fuller et al. (1999)

Spiny waterflea (Bythotrephes longimanus)

Muskoka lakes – Yan et al. (1992)

Kerfoot et al. (2011)

Spotted galaxis (Galaxias truttaceus) Australia – Lintermans (2004)

Steelcolor shiner (Cyprinella whipplei) Oklahoma, USA – Miller and Robison (1973), Fuller et al. (1999)

Stonecat (Noturus flavus) West Virginia, USA 1951 Fuller et al. (1999)

Striped shiner (Luxilis chrysocephalus) Alabama, Florida, West Virginia, USA


Suckermouth minnow (Phenacobius mirabilis)

Michigan, Ohio, New Mexico, USA

Great Lakes basin


Mills et al. (1993)

Swallowtail shiner (Notropis procne) New River, Virginia – Fuller et al. (1999)

Telescope shiner (Notropis telesopus) North Carolina, Virginia and West Virginia, USA

1940s-1950s Fuller et al. (1999)

Tennessee shiner (Notropios leuciodes) Chattahoochee River, Georgia

New River, Virginia

–

–



Threespine stickleback (Gasterosteus aculeatus)

Lake Huron

Mohave River, California

1980s

1938-40

Gibson (1982), Stedman and Bowen (1985) Fuller et al. (1999)

Tiger salamander (Ambystoma tigrinum) Arizona, California, Colorado, Louisiana, Nevada and Texas, USA

– Benson (1999)

Tonguetied minnow (Exoglossum laurae) West Virginia – Fuller et al. (1999)

Torrent sucker (Thoburnia rhothoeca) Virginia and West Virginia, USA 1950s Fuller et al. (1999)

Tui chub (Siphateles bicolor) Idaho, California and Nevada, USA – Fuller et al. (1999)

Utah chub (Gila atraria) Colorado, Idaho, Montana, Nevada, Oregon and Utah, USA


Virile crayfish (Oreconectes virilise) Seventeen U.S. states – Benson (1999)

Warpaint shiner (Luxilis coccogenis) North Carolina, South Carolina and Virginia, USA

1940s-1970s Fuller et al. (1999)

White perch (Morone americana) Several U.S. states – Feiner et al. (2013)

White River crayfish (Procambarus acutus) Six U.S. states – Benson (1999)

White shiner (Luxilus albeolus) North Carolina, Virginia and West Virginia, USA


White sucker (Catostomus commersoni) Colorado, Georgia, New Mexico, Utah, Wyoming, USA


Whitetail shiner (Cyprinella galactura) Kentucky, North Carolina, Virginia and West Virginia, USA

1980s Fuller et al. (1999)

Yellowfin shiner (Notropis lutipinnis) Georgia and North Carolina, USA – Fuller et al. (1999)


APPENDIX 3. Impacts of the introduction of aquatic invasive species and pathogens in a new waterbody (modified from Kerr and Grant 2000).

IMPACT REFERENCE

Interspecific Competition

Moyle (1976), Schoenherr (1981), Schaeffer and Margraf (1986), Evans and Loftus (1987), Fausch (1988), Kotler and Holt (1989), Parrish and Margraf (1990), Miller et al. (1992), Savino and Kolar (1996), Goodchild (1999), Dieterich et al. (2004), Carpenter (2005), Bergstrom and Mensinger (2009)

Predation Stenson (1972), Gorman (1974), Hurlbert and Mulla (1981), Post and Cucin (1984), Evans and Loftus (1987), Marsh and Brooks (1989), He and Kitchell (1990), Carvalho and Hauser (1995), Adams (1996), Marsh and Douglas (1997), Nystrom and Perez (1998), Ivantsoff (1999), Renai and Gherardi (2004), Correia et al. (2005), Peters and Lodge (2009)

Transmission of Disease and Parasites Dechtiar (1966), Hnath (1969), Summerfelt and Warner (1970), Stewart (1991), Robertson and Austin (1994), Goodchild (1999), Litvak and Mandrak (1999), Kiesecker et al. (2001), Perry et al. (2002), Sutherland et al. (2004), Tatem et al. (2006), Drake (2011), Goodwin (2012)

Genetic Impacts Ferguson (1990), Allendorf (1991), Arthington (1991), Krueger and May (1991), Vespoor and Hammar (1991), Gaffney and Allen (1992), Leary et al. (1995), Litvak and Mandrak (1999), Cross (2000), Rahel (2000), Scott and Helfman (2001)

Habitat Alteration King and Hunt (1957), Crivelli (1983), Lodge and Lorman (1987), Bauer (1988), Chambers et al. (1990), (Richardson and Whoriskey (1992), Bain (1993), Leach (1993), (Litvak and Mandrak (1993), Breukelaar et al. (1994), Roberts et al. (1995), Jude and Deboe (1996), Rhymer and Simberloff (1996), Aldridge et al. (2004), Rowe (2004), Rodriguez et al. (2005), Rowe (2007), Bakker and Dorenbosch (2013)

Community Alteration Evans and Loftus (1987), Fuller et al. (1999), Litvak and Mandrak (1999), Lodge et al. (2000), Yan et al. (2002), Wilson et al. (2004), Rodriguez et al. (2005), Burlakova et al. (2009), Wanink et al. (2010)

Economic Impacts Barbier (2001), Pimentel et al. (1999, 2000, 2005), Connelly et al. (2006), Lovel et al. (2006), Liu et al. (2011)

Displacement of Indigenous Species Deacon et al. (1964), Barlow et al. (1987), Nichols et al. (1990), Lassuy (1995), Taylor and Redmer (1996), Lodge et al. (2000), Kats and Ferrer (2003), Koel et al (2005), Gil-Sanchez and Alba-Tercedor (2002), Aldridge et al. (2004), DiStefano et al. (2009)

Adverse Effects on Recreational Fisheries Lodge et al. (2000)

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