Charles Jacoby Linda Walters Shirley Baker Karen Blyler

Florida Sea Grant College Program • http://www.flseagrant.org • SGEB 60

1 Acknowledgments Funding and support

We thank Mike Spranger for his encouragement This publication was supported by the National throughout the development of this primer. Sea Grant College Program of the U.S. Steve Kearl and Dorothy Zimmerman provided Department of Commerce’s National Oceanic valuable editorial input. and Atmospheric Administration (NOAA) under NOAA Grant No. NA16RG-2195. The views expressed are those of the authors and do not The authors necessarily reflect the views of these organizations. Charles Jacoby is an Assistant Professor and Extension Specialist in the Department of The Florida Sea Grant College Program, the Fisheries and Aquatic Sciences at the University Southeast Regional Aquatic Nuisance Species of Florida in Gainesville, Florida. (ANS) Education and Outreach Network, a partnership between the Sea Grant College Shirley Baker is an Assistant Professor in the Programs in Alabama, Florida, Louisiana, and Department of Fisheries and Aquatic Sciences at Mississippi, UF/IFAS Extension and the the University of Florida in Gainesville, Florida. University of Central Florida provided support for the development of this document. Linda Walters is an Assistant Professor in the Department of Biology at the University of Additional funds to support publication of this Central Florida in Orlando, Florida. document were provided by the Tampa Bay Estuary Program. Karen Blyler is the Coordinator of Education and Training in Aquatic and Environmental Science for the Florida 4-H Youth Development Program at the University of Florida in Gainesville, Florida.

Photo credits

Cover, upper left: Phyllorhiza punctata, an Indo-Pacific jellyfish now found in large numbers around the Gulf of Mexico. (Photo courtesy Dauphin Island Sea Lab)

Cover, center left: Perna viridis, an Indo-Pacific mussel that became established in Tampa Bay and has spread around Florida and into Georgia. (Photo courtesy U.S. Geological Survey)

Cover, lower left: Pterois volitans, an Indo-Pacific fish often kept in saltwater aquaria that is now found along the east coast of the United States. (Photo courtesy J.E. Randall)

Cover, bottom half: Aerial photograph of an explosion of the Australian spotted jellyfish (Phyllorhiza punctata) in the Gulf of Mexico. (Photo courtesy Dauphin Island Sea Lab)

Page 1, upper right: Pterois volitans, an Indo-Pacific fish often kept in saltwater aquaria, is now found along the east coast of the United States. (Photo courtesy J.E. Randall)

2 What are invasive species?

nvasive species have looking at the history of species I entered an area that is introductions, the harm caused by not part of their natural invasive species, how invasive range. They represent one type of species are introduced and become alien, non-native, nonindigenous invasive, and how we can manage or introduced species. In contrast, our own actions and activities to native or indigenous species occur help solve the problem. in an area naturally. Invasive species that are native or intro- How long have invasive duced differ from beneficial native species been around? and non-invasive species because they can or do cause harm to the People began introducing species as economy, the environment or soon as they could travel. For human health. Typically, invasions example, beginning around 1500, involve the spread of an introduced Europeans transported Old World species. Natural vectors, such as species to their new settlements in storms, or vectors associated with the Western Hemisphere and human activities may cause elsewhere. The manifests from Columbus’ second and subsequent People apply a confusing introductions and subsequent and muddled collection of invasions by moving species along voyages indicate deliberate trans- words to invasive species. pathways. Human introductions port of species regarded as poten- may be intentional or uninten- tial crops and livestock. A glossary found on pages tional, and both types of introduc- Humans continue to disperse 14 and 15 provides species, and the worldwide increase tions can cause bioinvasions. We definitions for key terms in plant, animal, and microbial can add a political bent to the used in this publication. discussion by applying the term introductions roughly tracks exotic to species that have entered the increase in human a new country and using the term transport and commerce. In transplanted for species moved to particular, the meteoric new areas within the same country. growth of global commerce Whatever definitions we choose, in the past 200 to 500 years we should all be concerned about has produced numerous invasive species and the harm they opportunities for biotic cause. First, we want to avoid invasions. During this time harmful consequences from span, the scope, frequency introductions. Second, our actions and impact of people’s or activities can spread invasive deliberate and accidental species, so we are responsible for movements of organisms managing both introductions and undoubtedly dwarf the any consequences. Managing effects of natural forces invasive species will not simply be during any 500-year period a matter of banning all introduc- in the earth’s history. For tions because all of us benefit from example, introductions to Figure 1. Rate of new invasions by marine invertebrates and seaweeds in the U.S. coastal some introductions and many coastal habitats in the United States have increased zone from 1790 to 1999 (374 total invasions). introductions are accidental or Source: Ruiz et al., 2000; Carlton, 2001. unintentional. We can improve our over 100-fold in the last management of invasive species by 200 years (Figure 1).

A PRIMER ON INVASIVE SPECIES IN COASTAL AND MARINE WATERS 1 Box 1. Throughout history, we have What harm do invasive Deliberate and accidental introductions of accidentally and deliberately intro- species cause? invasive species. duced numerous species. Our Introduced species typically do O deliberate introduction of few, if any, invasive tendency to accidentally or deliber- microorganisms, although we introduced ately introduce invasive species not survive and reproduce; there- yeasts for fermentation and mutualists, such varies among types of organisms fore, they seldom establish themselves as the symbiotic, mycorrhizal fungi found in (Box 1). The number of deliberately and become invasive (Figure 2). plant roots introduced species that become Introductions can be viewed as an O deliberate introduction of some insects that invasive emphasizes the fact that not ecological parallel to Russian have caused adverse consequences, such as all pests arrive unheralded and roulette. We ‘load bullets into a bumblebees in New Zealand, but the majority gun and spin its cylinder’ by of invasive insects have probably been inconspicuously. Many of our worst introduced accidentally problems are the product of our moving species beyond their own deliberate but flawed plans. natural ranges. Then, we ‘pull the O deliberate introduction of a few marine trigger’ by introducing species into invertebrates, such as the Pacific oyster imported from Japan to Washington state, but a growing number of invaders, such as the zebra mussel, have arrived accidentally as contaminants in ballast water and through other vectors O deliberately introduced most invasive vertebrates, principally fish, mammals and birds, although some of the worst vertebrate invaders have been spread accidentally, such as rats, brown tree snakes and sea lampreys O deliberately introduced many, if not most, plant invaders, including water hyacinth, melaleuca trees, salt cedar and other serious pests, although some invasive plants have been accidentally introduced as contami- nants among crop seeds and other cargo

Figure 2. Steps in becoming an invasive species.

2 A PRIMER ON INVASIVE SPECIES IN COASTAL AND MARINE WATERS new ranges and take our chances on efficiently. For example, Box 2. whether they cause harm by estab- invasive plants typically out Examples of how invasive species harm lishing and spreading. As in any compete natives by shading populations. game of chance, the outcome of a them or using scarce water. O Asian chestnut blight fungus arrived in New York City given round is uncertain. We may When a species interferes on nursery stock early in the 20th century and, within introduce numerous species or one with or harms another in the a few decades, the blight had spread and destroyed species numerous times and suffer no competition for resources, almost all the American chestnuts that had once dominated forests in the eastern United States significant consequences, or we may ecologists call it interference (disease) introduce a single species once and competition. Invasive species generate immense harm. In general, clearly demonstrate this tactic. O predatory Nile perch introduced into Africa’s Lake Victoria has eliminated or gravely threatens more species cause harm by affecting plant For example, many invasive than 200 of the 300 to 500 native species of small, and animal populations or commu- animals exhibit aggressive cichlid fishes (predation) nities, ecosystems and ecosystem behavior that drives native O feral and domestic cats introduced to every part of services, or our economy. animals out of their natural the world have depleted breeding populations of ranges, and invasive plants may seabirds and endemic land birds on oceanic islands Effects on populations produce toxins (phytotoxins) and contributed significantly to the extinction of that inhibit the success of six endemic birds in New Zealand and at least six Invasive species often affect species of Australian rodent-like marsupials populations or interbreeding groups native plants. (predation) Invasive species can also of organisms. They typically cause O North American gray squirrels replace the native extirpation or losses at the local level, eliminate native species by red squirrels in Britain because they forage more but they possess the potential to hybridizing with them. Such efficiently (scramble competition) interbreeding is a particular cause extinction or losses at the O succulent ice plant, Carpobrotus edulis, forms a mat global level. Extinction becomes a danger when the native popu- that shades native plants in coastal California and very real possibility for endangered, lations are endangered, threat- removes scarce water that the native species need threatened or rare species. Invasive ened or rare. Hybridization (scramble competition) species typically affect native popula- between a nonindigenous O introduced ant species – the red fire ant, the tions through six types of ecological species and a native one can Argentine ant and the big-headed ant – all interactions (Box 2). even produce a new invasive significantly reduce native ant populations by aggression (interference competition) Invasions by pathogens can species. In addition, hybridiza- severely impact native species. For tion can threaten a native O Quackgrass damages crops, and it may do so by producing phytotoxins (interference competition) example, we all know of plant species even when the hybrids diseases that have spread readily, die, simply because crossbreed- O hybridization with the introduced North American especially those that affect crops, ing reduces the number of new mallard threatens the genetically distinct existence offspring added to the species’ of both the New Zealand gray duck and the Hawaiian livestock or pets. We also could duck (interbreeding) include effects on human health own population. North American cordgrass, carried in shipping ballast here, such as concerns about ballast Finally, invasive species can O evolve new ways to threaten to southern England, hybridized with native water introducing cholera in places it cordgrass and one hybrid underwent a genetic is not normally found. natives after establishment. change to produce a fertile, highly invasive species Invaders can directly remove Plants and animals exhibit of cordgrass (interbreeding) remarkable capacities to native species by predation and O females of the European mink, already gravely grazing. Historically, people have develop resistance or other threatened by habitat deterioration, hybridize with introduced many animals in ways to overcome the defenses males of introduced North American mink resulting attempts to make new places more erected by native species. in aborted embryos and a loss of eggs that exacerbates their decline (interbreeding) like ‘home.’ Even today, we continue to release domesticated animals into Effects on communities, O parasitic wasps imported to the United States to ecosystems and ecosystem control the alfalfa weevil were originally ineffective new places where they can cause against the Egyptian alfalfa weevil, but, fifteen years significant harm. services later, the wasps appeared to have evolved a Nonindigenous species also harm In the direst situations, resistance because 95% of their eggs were surviving natives by competing for scarce invasive species threaten to in the larvae of the Egyptian alfalfa weevil (evolution) resources in what ecologists call disrupt both communities, scramble competition. Invasive which are formed by groups of species often seem capable of out populations, and entire ecosys- competing native species by seques- tems, which are natural sys- tering water, nutrients or food more tems that produce and recycle

A PRIMER ON INVASIVE SPECIES IN COASTAL AND MARINE WATERS 33 Box 3. more energy and other resources than Effects on economies Examples of how invasive species harm they import (Box 3). Invasive plants Public and governmental support communities, ecosystems and ecosystem demonstrate these threats best. Typi- for the prevention or control of inva- services. cally, invasive plants change ecosys- sions often falters because most people O Australian paperbark or melaleuca trees tems by using significant amounts of don’t understand or appreciate the replaced cypress, sawgrass and other water, occupying space and shading complex links between nature and our south Florida natives at the rate of natives, providing poor habitat and 20 hectares per day resulting in about economies. Invasive species pose 160,000 hectares that provides poor forage for native animals, or altering threats to biodiversity and processes in habitat for many native animals, uses fire regimes. In fact, many invasive ecosystems, and these threats translate huge amounts of water, and intensifies plants wreak havoc on ecosystems by directly into economic consequences, fire regimes fostering more frequent or intense fires such as losses of crops, forests, fisheries O a vine-like perennial shrub from South that they withstand better than key and grazing lands. However, the America, Chromolaena odorata or Siam native species. The paperbark tree, a economic consequences of biotic weed, aggressively suppresses Melaleuca species, has this effect in regeneration of primary forest in both invasions are poorly explored and Asia and Africa and provides feeding Florida, as do numerous invasive quantified. Although we have anec- niches that sustain other pests grasses worldwide. Grasses, in particu- dotal evidence of local and even lar, produce a great deal of flammable O a highly invasive neotropical shrub, regional costs from invasive species, we Lantana camara, increases the standing dead material, dry out lack clear, comprehensive information occurrence of sleeping sickness among rapidly, and resprout quickly after on these costs at national and global animals and humans by serving as fires. levels. additional habitat for the tsetse fly that Invasive plants may also alter Biotic invasions cause three main normally lives along East African ecosystems in more insidious ways. For streams types of economic impact. First, example, they may change the cycling invasive species reduce potential O a small tree native to the Canary Islands, of nutrients in a way that encourages Myrica faya, is transforming the entire economic output by reducing the ecosystem in Hawaii Volcanoes invasions by other plants, or they may production, survival and fitness of National Park by fixing nitrogen and promote invasions by non-native crops, domesticated animals and causing a 90-fold increase in supplies animals. In some cases, alien plants fisheries. Second, we suffer direct costs of this scarce nutrient, which encourages and animals combine to create a associated with combating invasions, many other introduced plants to enter positive feedback loop with the plant the ecosystem including all forms of quarantine, providing food for the animal and the control and eradication. Combating O Myrica also attracts the introduced animal distributing the plant’s seeds. Japanese white-eye, the most invasive species that are threats to destructive bird invader in native In some places, invaders cause human health, either as direct agents Hawaiian forests, and the white-eye ecosystem changes that profoundly of disease or as vectors or carriers of exacerbates ecosystem changes by alter the landscape and the ecosystem disease-causing parasites, represents a dispersing Myrica seeds services we value. For example, blue- third category of costs. O the Bluegrass Country of Kentucky grass in Kentucky and some other These costs form a hidden but invokes images of a pastoral or pristine ecosystems we take for granted in the onerous ‘tax’ on many goods and setting; however, bluegrass is a Eurasian United States are actually a result of services. Tallying these costs represents invader introduced during European successful invasions by species intro- settlement and land clearing after which a formidable task. One group recently it replaced the original open forest, duced by colonists. Overall, our attempted to tabulate the annual cost savanna, and understory of wild rye and knowledge of invasions is poor, but of all nonindigenous species in the possibly canes known invasions raise the concern that United States. They estimated that O burning of forests and introduction of the current pace and extent of inva- nonindigenous weeds that affect crops African grasses to the Amazon basin sions may significantly influence other cost agriculture about $27 billion per creates grasslands that contain much agents of global change in an unpre- year, based on a potential crop value of less plant biomass and sequester less dictable manner. For example, we may carbon, which could exacerbate the $267 billion. Loss of forage and the buildup of carbon dioxide in the alter the global carbon cycle by application of herbicides to weeds in atmosphere and influence the global converting diverse forests to grasslands. rangelands, pastures and lawns cost a climate Grasses create less tissue or biomass so further $6 billion each year. When the they use and sequester less carbon group combined these types of direct dioxide. Less carbon in plants means losses with indirect costs for activities more carbon dioxide in the atmo- such as quarantine, the total cost of all sphere, which may accelerate global nonindigenous species (plants, animals warming and the attendant conse- and microbes) exceeded $138 billion quences.

4 A PRIMER ON INVASIVE SPECIES IN COASTAL AND MARINE WATERS per year. By any standard, such costs example, we are uncertain about the Box 4. are formidable, even for a productive, origins of ‘killer dinoflagellates’ in the Facts about marine invasions. industrialized society such as the genus Pfiesteria and many toxic algal O a tropical seaweed, Caulerpa taxifolia, United States. blooms or red tides. Overall, informa- developed tolerance for colder These estimates illustrate the tion indicates that coastal and marine temperatures while growing in public and anecdotal and preliminary nature of introductions have vast negative effects private aquaria in Europe, and, when it escaped into the northwest Mediterranean, our current understanding of the on the economy of the United States. its new tolerance of winter temperatures economics of invasions. One solution In addition, funds diverted to research helped it blanket large stretches of the would be a more frequent application and control of introduced species seafloor and threaten nearshore plant of economic tools such as cost:benefit could have been used for other pur- populations analysis when considering proposals to poses. Between 1999 and 2001, the O the European periwinkle, introduced to Nova Scotia around 1840, grazes heavily on import species for perceived economic costs for research and control related to shoreline plants that trap and hold benefit. When it comes to future three Pacific coast introductions sediments. As a result, many coastal inlets movements of species, society needs to equaled nearly one-third of all the transformed from mudflats and salt consider results from analyses like funds available through the Sea Grant marshes to rocky shores those provided for hydroelectric dams, College Program for an entire decade O introduced species foul jetties, marinas and canals, airports, and other projects of research and education on intro- buoys with the result being changes in food webs and energy flow that further stress with potential environmental conse- duced species. troubled fisheries quences. Scientists predict that Although poorly documented, O introduced species of crabs, mussels, clams, cost:benefit analysis of many deliber- introduced marine species fundamen- jellyfish, seagrasses and marsh grasses form ately introduced invaders would tally alter the biodiversity of coasts in conspicuous parts of marine ecosystems demonstrate forcefully that their costs the United States and around the from the Hawaiian Islands to the Pacific Northwest, south to San Francisco Bay and to society swamp any realized or world. Invasive species often out southern California, east to the Gulf of perceived benefits. compete native species either in a Mexico, and north to Chesapeake Bay and scramble for resources or through New England Those are terrestrial and interference competition. In some O Puget Sound, in Washington State, Coos Bay, ecosystems, the introduced species in Oregon, and Chesapeake Bay, in Virginia freshwater examples. and Maryland, harbor at least 50, 60 and become so dominant that finding 43 non-natives respectively What about impacts on native species becomes an elusive goal. O more than 175 species of introduced marine coastal and marine waters? Overall, the effects of introduced invertebrates, fish, algae and higher plants The economic and ecological effects species rank second to habitat destruc- live in San Francisco Bay alone of coastal and marine introductions are tion as a threat to biological diversity. O in San Francisco Bay alone, an average of notoriously difficult to estimate, and Communities and ecosystems are also one new introduction becomes established every 14 weeks between 1961 and 1995 detailed estimates have not been made affected by introductions of new predators, competitors, disturbers, O new introductions and establishments occur for the waters of the United States. In in the United States in 2000–2001, with: any estimate, negative and positive parasites and diseases. These introduc- tions lead to vast alterations in species vast numbers of Pacific spotted jellyfish effects must be considered. Negative (Phyllorhiza punctata) invading the Gulf of effects include revenues lost due to the interactions and to changes in nutrient Mexico cycling and energy flow that result in destruction of fisheries; loss of other billions of small carnivorous, European resources due to predation, competi- cascading and unpredictable effects flatworms (Convoluta convoluta) appearing tion, or disease; the cost of removing throughout entire communities. in the Gulf of Maine introduced fouling organisms from Although no one has determined the the Asian whelk (Rapana venosa) becoming hundreds of thousands of recreational actual economic impact, certain more abundant in Chesapeake Bay vessels; and the costs associated with indicators suggest that species intro- the Japanese mahogany clam (Nuttallia the damage to pilings and other duced into coastal and marine systems obscurata) reaching southern Oregon structures by introduced wood-borers. may cost the United States hundreds the brown mussel (Perna perna) invading the Gulf of Mexico Positive effects include the value of of millions of dollars every year. Although our knowledge of impacts thousands of Asian shore crabs fisheries based on non-native species (Hemigrapsus sanguineus) in Long Island and the aesthetic value of introduced from introductions is poor, we now Sound know that numerous species have been species, even if the public does not the Mediterranean green seaweed know that the species are introduced. or are being introduced to the coastal (Caulerpa taxifolia) and the Asian kelp A further challenge is estimating the and marine ecosystems of the United (Undaria pinnatifida) appearing in potential effects of cryptogenic species States (Box 4). Hundreds of intro- southern California whose origins are uncertain. For duced species occur in the United thousands of farmed Atlantic salmon (Salmo salar) escaping into the Pacific

A PRIMER ON INVASIVE SPECIES IN COASTAL AND MARINE WATERS 5 Box 5. States’ waters, and the rate of known vessels ply the world’s seas. These How many and what kinds of species occur introductions has increased exponen- vectors transport a diverse array of in ballast water? tially since the 18th century, with no marine life at unprecedented rates O studies in the United States, Germany, signs of leveling off. Unfortunately, the (Box 5). In a variety of ways, these Scotland, Wales, Australia and Hong exact number of coastal and marine vessels carry living aquatic organisms Kong, reveal that ballast water can introductions remains unknown for a from fresh, brackish or marine water, contain representatives of all major variety of reasons. For example, we across and between oceans, or along and most of the minor groups of marine life seldom know the origin and history of coastlines. An arriving vessel may be a coastal or marine species. Scientists virtual ‘floating biological island,’ with O many species travel as larval, or dispersal, probably overlook introductions of hundreds of species living both on and stages, and then they become bottom- dwelling organisms as adults both microscopic species and groups of in the ship. organisms that are difficult to identify. Since the 1880s, most ships carry O examples of commonly transported species include sea anemones, worms, Some introduced species are cryptoge- ballast water. Ballast water pumped or barnacles, crabs, snails, clams, mussels, nic, which means they look similar to fed by gravity into a vessel compen- oysters, bryozoans, sea urchins, sea native species and can only be distin- sates for the lack of cargo, maintains squirts and algae guished by genetic analyses. A decline the trim and draft of the vessel, and O other species, such as diatoms, dino- in coastal and marine exploration and enhances stability and operational flagellates, copepods and jellyfish, reduced training in systematics and efficiency. Most ships, even those with travel as adults in ballast water taxonomy contribute to confusion cargo, carry some ballast water. A ship O certain viruses and the bacteria that about the precise number of marine may take on ballast water in port or at cause cholera epidemics have also been introductions. Overall, we seem sea. The water is held in ballast tanks detected in ballast water determined to play ecological roulette or floodable cargo holds. A ship may O overall, organisms in ballast water with introductions of marine species. discharge all or some of its ballast range in size from microscopic viruses water when it arrives at its next port. and bacteria to fish 12 inches (30 cm) or longer How are species introduced Before water became commonly used into coastal and marine as ballast, ships were loaded with O we probably transport at least rocks, sand, soil or almost anything 7,000 different species of marine life in systems? ballast water each day cheap and heavy. Movement of ‘dry Every day, a large number of human- ballast’ spread thousands of species of O ballast water, carrying all these non- mediated vectors move thousands of native species arrives in the United insects and other arthropods, mollusks States at the rate of 2 million gallons coastal or marine organisms around the and plants. In contrast, ballast water per hour world and introduce them into new tends to spread saltwater species from areas. Recent events underscore the four communities: need for effective education and man- Š plankton: organisms that drift agement programs that increase aware- passively or swim modestly; ness of how our activities result in Š nekton: species that swim freely; introductions. Species arrive in United Š fouling: organisms (including States’ waters from virtually every bacterial films) that attach to the region of the world after traveling along vertical walls and horizontal numerous trade routes that continu- structures in the ballast ously change. At a given coastal site in compartments; and the United States, our activities and Š benthos: organisms that dwell on actions create the potential for numer- the bottom, such as beds of marine ous, repeated and frequent introduc- worms and associated species and tions of non-native species. An exami- the encysted, or resting, stages of nation of some of the key vectors plant plankton (phytoplankton) and illustrates the extent and depth of the animal plankton (zooplankton). challenges we face. Ballast is not the only means by which ships carry marine life from Ballast water and fouling foreign shores to the United States. organisms from ships Fouling organisms attached to the Today, more than 45,000 commer- outside of vessels can also invade new cial, cargo-carrying vessels and hun- regions. For the past 500 years, tens of dreds of thousands of recreational thousands of vessels have formed a

6 A PRIMER ON INVASIVE SPECIES IN COASTAL AND MARINE WATERS conveyor belt between North America other resources generate a concomitant Box 6. and the rest of the world by transport- increase in the movement of dry docks Examples of introductions related to ing species that could not survive used to float and repair ships and fisheries activities. drifting on their own across the high semi-submersible, self-propelled O intentional introduction of Atlantic seas. Historically, barnacles, mussels, drilling and production platforms striped bass (Morone saxatilis) to the hydroids, algae and other marine used for discovery and extraction of Pacific coast in the 1870s represents a organisms formed fouling assemblages resources. These structures have striking example of creating a new fishery thick enough to harbor free-living abundant subsurface space for fouling species, such as crabs and fish. Modern communities and ballast water O live Atlantic lobsters (Homarus ships continue to carry fouling organ- systems. americanus) purchased at the airport in Boston or New York are released in isms on their hulls, rudders, propellers Other maritime activities can also southern California waters hours later and propeller shafts, as well as in transport and introduce species. by people who would prefer to let the seawater piping systems (including Examples include the long-distance shellfish live rather than eat them ballast intake screens) and their sea movement of navigation buoys, marina O seaweed used to pack bait-worms from chests, the water compartments floats, amphibious vessels and sea- the Atlantic coast apparently leads to between the outside of ships and the planes. the introduction of the European shore ballast pumps. Sea chests, in particu- crab (Carcinus maenas) on the Pacific coast lar, accumulate organisms that would Fisheries activities not survive on the hull of the ship, and O live and healthy softshell, blue crabs Marine life is dispersed through a (Callinectes sapidus) from Chesapeake they are now suspected of playing a wide range of fisheries activities Bay have been bought in San Francisco significant role in introductions. (Box 6). These activities fall into two fish markets and occasionally released Antifouling paints, which are toxic to broad categories: intentional releases, into California waters by the public marine life, discourage or prevent the whether legal or illegal, and accidental O proposals to revitalize the waning attachment of fouling organisms. releases. oyster industries in the Gulf of Maine Regulations that protect the environ- The wisdom of deliberately intro- and Chesapeake Bay have focused on the use of non-native species with ment by decreasing use of certain ducing non-native species has been paints may increase the presence of resistance to the diseases that killed debated for a long time. In the native Chesapeake Bay oysters fouling populations. For example, 19th century, many ‘acclimatization paints containing tributyl tin (TBT) O Virginia placed sterile stocks of the societies’ and government agencies Japanese oyster (Crassostrea ariakensis) probably will be banned internation- intentionally released non-native in Chesapeake Bay in 2001 ally. In Australia, decreased use of species into United States’ waters to paints with TBT has already increased O Maryland did not participate in the ‘improve nature’ or start new fisheries. stocking experiment because of ship fouling. Further studies are Today, we make far fewer purposeful potential liabilities if some individuals needed to determine if there is a attempts to sprinkle exotic marine revert to a reproductive state, establish correlation between decreased use of species into the wild. Current propos- wild populations and spread to states TBT-based paints and increased and jurisdictions that do not want non- als to introduce species often focus on indigenous oysters introductions from ship fouling. using individuals that cannot repro- Both ballast water and ship fouling duce because they are sterile or because O this incident points out the difficulties may combine to introduce some of managing introduced species under temperatures and other conditions for the current regulatory system species of marine life. Ballast water and reproduction are not right. Concerns the organisms in it are introduced remain because individuals could when they are discharged as a ship adapt to new temperature regimes and loads cargo. Organisms attached to other conditions, and individuals may ships’ hulls or sea chests must repro- not remain sterile. Regardless, people duce, become dislodged or swim off still deliberately transport an unknown the ship in order to be introduced. number of species and individuals on a Many organisms, including barnacles, daily basis. These animals are trans- mussels, hydroids, sea squirts and algae, ported for direct consumption as live can be introduced by both mechanisms. seafood, for use as live bait with ‘freshening’ in the marine environ- Dry docks, drilling platforms and ment, or with the intent or hope of maritime activities starting a new fishery. We have no data Increases in international commerce on the scale of illegal attempts to start and reliance on offshore oil, gas and new fisheries by introducing species.

A PRIMER ON INVASIVE SPECIES IN COASTAL AND MARINE WATERS 7 Such attempts may be a significant tropical fish represent an abundant source of introductions, given the and diverse component of fish ease with which people bring living communities. Data indicate that organisms into the United States in accidental or deliberate releases of their luggage. aquarium species may contribute The distinction between inten- to this situation (Figure 3). tional and unintentional introduc- In addition to marine life tions becomes blurred when we available for home aquaria, a long include innumerable incidents in list of other marine organisms can which the public releases living, be purchased for education and nonindigenous organisms without research on scores of websites. The any particular future intent. After a fate of these organisms remains day’s fishing, fishermen may discard largely in the hands and control of leftover bait-worms and the seaweed the public, educators or they were packed in, which contains researchers with few, if any, numerous other live organisms that regulatory constraints and little or can become invasive. As a vector, no information about the potential fishermen move relatively small consequences of accidental or amounts of material and few species, deliberate releases. but they demonstrate that a small vector may not be a minor one if it Other vectors leads to major introductions. This A variety of additional vectors, situation raises the challenge of such as disposal of dredge spoil, whether to prioritize management beach renourishment and according to the volume of traffic movement of recreational through a vector or the number of equipment, have been invoked at species that are transported. The one time or another to explain the sale of live seafood in markets where appearance of new species. The it is not native represents another significance of these vectors may troublesome vector. Laws may vary regionally or locally, with require private persons or public bursts of activity related to special agencies to obtain permits before or focused events. Biocontrol, or releasing animals to deliberately the release of one species to control start a fishery, but seafood consum- another, has not been tried in the ers can release animals without oceans, but it is increasingly a being aware of or affected by such topic of discussion. requirements. Floating marine debris as a potential vector for introductions Aquarium industry and the represents another area of availability of living marine increasing concern, especially in organisms on the Internet the Pacific Ocean. Large numbers In the United States, the of fishing nets drift in the Pacific aquarium industry sells an unknown and come ashore covered with number of invertebrates, fish, algae marine organisms in places like the and seagrasses as ‘pets’ or ‘displays.’ Hawaiian Islands. Tropical and subtropical species form the bulk of the imports, but Coastal dispersal of temperate species are also traded. A introductions few species are regulated, such as Once a new species has arrived piranhas, but most imports are not and become established in the restricted. Although the organisms United States, it may disperse are not imported for release into the along the coastline by natural wild, in the southern half of the means. It may float or drift as a United States, subtropical and planktonic larva, or it can be

8 A PRIMER ON INVASIVE SPECIES IN COASTAL AND MARINE WATERS Figure 3. Sources of introductions in Florida. The large contribution by aquarium releases is unusual. Source: USGS, 2001. attached to floating materials such used. Crabs were moved as bait, as as seaweeds, seagrasses, and marine aquarium species, as part of the debris. Human-mediated vectors school-educational market, as can also transport a new species research animals, accidentally with alone or in combination with lobster shipments, and along with natural means. petroleum production platforms. Most research efforts focus on The stage for new invasions is mechanisms that could bring new, constantly set and reset because exotic species into United States’ numerous transport mechanisms waters. As a result, few data ad- come into play on an hourly basis. equately describe or quantify how Numerous ships and seaplanes new invaders spread along a coast- reach coastal sites in the United line. Such data are critical to States. Marina floats drift in from predict both the rate and direction distant harbors. Live, non-native, of spread. marine organisms are purchased for food, for use as bait or as pets. The number and diversity of Educational institutions, research transport vectors institutions and public aquaria In the past 200 years, the hold a large variety of living, exotic, number of vectors transporting marine organisms. A large seafood marine species has steadily restaurant overlooking the water increased. In 1800, only two imports live Maine lobsters, mechanisms, ship-hull fouling and wrapped in fresh, invertebrate- ballast rocks, were available to move laden seaweed. In a nearby salt the European shore crab (Carcinus marsh or estuary, restoration of maenas) across or between oceans. important plants may be By 1900, three additional underway, using stocks and mechanisms became available: sediments shipped from distant ballast water, intentional movement locations. Today, more species and as food, and the importation of more individuals of those species oysters for aquaculture. By the are transported because of the year 2000, ten human-mediated increasing diversity of vectors. mechanisms could move the crab Moreover, the fact that most around the world. Six new vectors species can now be introduced by were added to the previous vectors many different vectors makes the with ballast rocks being no longer prevention of introductions an even greater challenge.

A PRIMER ON INVASIVE SPECIES IN COASTAL AND MARINE WATERS 9 Box 7. Why can’t we just exclude Who’s managing all of this? Generalized characteristics of successful potential invaders? Management of invasive species invasive species. Management of invasive species occurs at the international, national O high rate of reproduction would be far simpler if we could and state levels. Internationally, the O colonizing disturbed habitats before identify potential invaders and where focus has been on ballast water, other species they might become established before endangered species, agricultural pests O short generation time we have a problem. Scientists have and human health (see Appendix A). listed characteristics of both potential O long life span Terrestrial and freshwater species invaders (Box 7) and places susceptible remain the dominant concerns at all high dispersal rate O to invasion (Box 8), but using these levels, but concerns about saltwater O single-parent reproduction by a female generic lists to manage invasive species species are increasing. that has eggs or is pregnant proves to be difficult. Within the United States, invasive O asexual reproduction Applying the generic lists proves to species management is being addressed be difficult because: O high genetic variability by coordinated efforts across multiple Š species may only display one of the agencies (Figure 4). The coordination O phenotypic plasticity, which means it typical characteristics; is to be conducted through the Inva- changes form in response to environ- Š many species do not display any of mental conditions sive Species Council, a national man- the characteristics in their native agement plan and the Invasive Species O broad native range habitat; Advisory Committee (Figure 5). abundant in native range Š invasions may involve a particularly O Further coordination regarding aquatic invasive species, a particularly O tolerant of a wide range of conditions nuisance species and linking to states vulnerable habitat, or both; and and regions occurs through the O habitat generalist, which means it lives Š the causes of most invasions are in a variety of habitats Aquatic Nuisance Species Task Force unknown because they were noticed and its regional panels. In addition, O broad diet only after they occurred. O gregarious, which means it lives in groups O benefits from living with humans without causing us harm, that is lives as a commensal

Box 8. Generalized characteristics of habitats susceptible to invasion. O climatically similar to the original habitat of invader O recently disturbed by humans or natural events O low diversity of native species O absence of predators that will eat the invading species O absence of native species similar in ecology or morphology (form or structure) to the invader O absence of predators or grazers similar to the invader so food sources are naive O absence of fire so the invader will not need to adapt to this disturbance O simple food web with few connections so the invader will not be competing Source: Invasive Species: Federal and Selected State Funding to Address Harmful Nonnative Species, with many other species GAO/RCED-00-219 (Washington, D.C.: Aug. 24, 2000). Figure 4. Key Federal departments and their responsibilities for invasive species.

10 A PRIMER ON INVASIVE SPECIES IN COASTAL AND MARINE WATERS Source: National Invasive Species Council

Figure 5. Formation and roles of the Invasive Species Council and Invasive Species Advisory Committee.

A PRIMER ON INVASIVE SPECIES IN COASTAL AND MARINE WATERS 11 Box 9. states are appointing people to control of introductions as the Incidences that illustrate a need for effective deal with invasive species and key to success. Once a species vector awareness and management tools. developing management plans in establishes and spreads, 1989 seaweed with bait concert with national efforts (see eradication or control become far Appendix B). more costly and less likely. Seaweed used as packing for bait worms from Maine is discarded in Management plans at all levels Surveillance for nonindigenous San Francisco Bay. As a result, the typically recognize prevention or species and rapid removal before carnivorous European shore crab they establish and spread is the (Carcinus maenas) and the Atlantic second best defense. Management rocky shore snail (Littorina saxatilis) plans also recognize the need for invade the Pacific Coast. Box 10. research, education and outreach. 1992 illegal live imports What can I do about introduced species in Gathering and disseminating coastal and marine systems? The Chinese mitten crab (Eriocheir better information can help us sinensis) is discovered in San You can: manage invasive species. For Francisco Bay. Illegally imported live O never release aquarium species, bait, example, informed decision- animals from Asia are intercepted fish, water garden plants or other plants at California airports. makers should make better and animals into the wild choices about large-scale, 1998 historic battleship O remove attached plants and animals intentional introductions, and an To soak the hull in freshwater, the from boats, trailers, SCUBA gear, anchors informed public can modify their and all other equipment USS Missouri is moved into the behavior to prevent small-scale, Columbia River prior to heading for O drain water from boats, live wells and the Hawaiian Islands. However, the intentional or accidental bilges before entering new areas lower hull remains in saltwater, and introductions. as a result, the Mediterranean mussel O only release ballast water as directed Along with others, the (Mytilus galloprovincialis) establishes by the United States Coast Guard National Sea Grant College in Pearl Harbor and colonizes the O never use non-native species for bait ballast tanks of a submarine. Program and its state programs contribute to management efforts 1998 mariculture (see Appendix C). For example, Atlantic salmon (Salmo salar) escape the National Sea Grant College from farms in the Pacific Northwest. Program awards grants for 2000 marina floats research and outreach related to Floats are towed at sea from New invasive species. Partnerships Jersey to Massachusetts even among other Sea Grant College though they are encrusted with Programs, such as Alabama, fouling organisms, including the Florida, Louisiana and Asian Crab (Hemigrapsus sanguineus). Mississippi, coordinate efforts 2000 home aquarium dealing with invasive species in The Mediterranean green seaweed their coastal and marine systems. (Caulerpa taxifolia) is apparently Key projects include workshops released from a home aquarium to help formal and non-formal into a lagoon near San Diego. It creates a well-established popu- educators incorporate invasive lation and eradication efforts follow. species into their classroom activities. 2000 live seafood Certainly, we can find times A hotel sushi bar releases live, where our management of coastal Japanese freshwater crabs (Geothel- phusa dehaani) into Lake Las Vegas and marine invasive species fails in Nevada, and they are later found (Box 9). All of us can help walking around. improve future efforts (Box 10). 2001 raw shellfish New Jersey embargoes 6,000 cases of raw clams from China, which were labeled ‘cooked.’ Hepatitis A virus is found in the shellfish.

12 A PRIMER ON INVASIVE SPECIES IN COASTAL AND MARINE WATERS Bibliography

Benson, A.J., P.L. Fuller and C.C. Jacono. 2001. Summary report of nonindigenous aquatic species in U.S. Fish and Wildlife Service Region 4. United States Geological Survey, Gainesville, Florida. 68 pp. Benson, A.J., D.C. Marelli, M.E. Frischer, J.M. Danforth and J.D. Williams. 2001. Establishment of the green mussel, Perna veridis (Linnaeus 1758) (Mollusca: Mytilidae) on the west coast of Florida. Journal of Shellfish Research 20: 21–29. Carlton, J.T. 2001. Introduced species in U.S. coastal waters: environmental impacts and management priorities. Pew Oceans Commission, Arlington, Virginia. 29 pp. Carlton, J.T. and J.B. Geller. 1993. Ecological roulette: the global transport of nonindigenous marine organisms. Science 261: 78–82. Florida Invasive Species Working Group. 2002. Statewide invasive species strategic plan for Florida 2002. Florida Invasive Species Working Group, Tallahassee, Florida. 44 pp. Gulf of Mexico Regional Panel. 2001. Gulf of Mexico Regional Panel: Aquatic Nuisance Species Annual Report 2001. Gulf of Mexico Program, Stennis Space Center, Mississippi. 22 pp. Mack, R.N., D. Simberloff, W.M. Lonsdale, H. Evans, M. Clout and F. Bazzaz. 2000. Biotic invasions: causes, epidemiology, global consequences and control. Issues in Ecology 5: 1–21. Meinesz, A., J. de Vaugelas, B. Hesse and X. Mari. 1993. Spread of the introduced tropical green alga Caulerpa taxifolia in northern Mediterranean waters. Journal of Applied Phycology 5: 141–147. National Invasive Species Council. 2001. Meeting the invasive species challenge. National Invasive Species Council, Washington, D.C. 74 pp. Pederson, J. (editor). 1999. Marine bioinvasions: proceedings of the First National Conference. Massachusetts Institute of Technology Sea Grant College Program, Cambridge, Massachusetts. 427 pp. Ruiz, G.M., P.W. Fofonoff, J.T. Carlton, M.J. Wonham and A.H. Hines. 2000. Invasion of coastal marine communities in North America: apparent patterns, processes, and biases. Annual Review of Ecology and Systematics 31: 481–531. Shafland, P.L. and W.M. Lewis. 1984. Terminology associated with introduced organisms. Fisheries 9: 17–18. United States General Accounting Office. 2002. Invasive species: clearer focus and greater commitment needed to effectively manage the problem. United States General Accounting Office, Washington, D.C. 103 pp. Williams, J.D. and G.K. Meffe. 1998. Nonindigenous species. In: Mac, M. J., P. A. Opler, C. E. Puckett Haecker, and P. D. Doran. Status and trends of the nation’s biological resources. 2 vols. United States Department of the Interior and United States Geological Survey, Reston, Virginia. 570 pp.

A PRIMER ON INVASIVE SPECIES IN COASTAL AND MARINE WATERS 13 Glossary

alien species. Any species that is not native to a given ecosystem, including seeds, eggs, spores, or other material capable of propagating the species. ballast water. Water used to adjust the trim and stability of a vessel. biocontrol. The release of one species in an effort to control another. bioinvasions. Introductions assisted by humans and natural range expansions. community. A group of populations that occupy the same area and interact. cryptogenic species. A species that may be introduced but we cannot distinguish it from similar natural populations without genetic analyses. ecosystem. Natural systems that produce and recycle more energy and other resources than they import. ecosystem services. Things or activities that we value and would not have without a healthy ecosystem to support them. endangered species. A species under threat of imminent extinction. established. A nonindigenous species that produces self-sustaining populations or a species that is naturalized. exotic species. A species that has been introduced into a country that does not contain any of its natural range. extinction. The complete global disappearance of a species. extirpation. The local disappearance of a species, as opposed to extinction. fouling organism. Animals and plants, such as barnacles, mussels and seaweeds, that attach to human-made substrates, such as piers, navigation buoys and the bottoms of ships. grazing. Feeding on plants. hybridize. To interbreed. indigenous species. A species that has not been introduced but rather historically occurred or currently occurs in an ecosystem. intentional introduction. The purposeful import, introduction or transplant of a nonindigenous species into an area or ecosystem where it was not previously established. interference competition. Interactions during the pursuit of resources, such as space, food or water, in which one species interferes with or harms another. introduced invasive species. Species that undergo population explosions and cause harm after being transported to a new area by human actions or activities. introduced species. A species whose presence in a given region can be traced to human actions or activities that dispersed it across natural geographic barriers or produced conditions favorable to its growth and spread.

14 A PRIMER ON IINVASIVE SSPECIES IN CCOASTAL AND MMARINE WWATERS introduction. The movement of an animal to an area outside its natural range through human activities. invasion. The spread of a species to a new area due to natural causes or human introduction. invasive species. A species that causes or is likely to cause harm to the economy, environment or human health. native species. A synonym for indigenous species. native invasive species. Species that get into modified habitats by their own means and then go through population explosions and cause harm. naturalized. A synonym for established. natural range. The area occupied by a species other than where it has been introduced. nonindigenous species. A synonym for alien species. non-invasive species. A nonindigenous species that does not spread but remains localized within its new environment. non-native species. A synonym for alien species. pathogen. A specific agent that causes a diseases, such as a bacterium or virus. pathway. The route or geographic corridor for an invasion. pest. Can be treated as a synonym for invasive species. population. An interbreeding group of organisms. predation. Feeding on animals. scramble competition. Interactions during the pursuit of resources, such as space, food or water, in which one species captures the resources without interfering or harming another. threatened species. Any species that is likely to become endangered within the foreseeable future throughout all or a significant portion of its range. translocation. A synonym for introduction. transplanted species. A species moved from its natural range to another area within the same country. unintentional introduction. An introduction resulting from activities other than the purposeful or intentional movement of a species, including transport in ballast water or in water used to transport plants or animals for aquaculture or other purposes. vector. A method of introduction.

A PRIMER ON INVASIVE SPECIES IN COASTAL AND MARINE WATERS 15 Appendix A. International laws and conventions related to invasive species.

Agreement Relevance Web site

International Plant Protection prevent introduction and spread of http://www.fao.org/legal/treaties/004t-e.htm Convention (IPPC) pests affecting plants

Agreed Measures for the require permits for introduction of http://www.antarctica.ac.uk/About_Antarctica/Treaty/ Conservation of Antarctic Fauna nonindigenous plants and animals Flora_and_Fauna.html and Flora into the Antarctic Treaty Area

The Convention on Wetlands protect wetlands from various http://www.ramsar.org/index.html (Ramsar Convention) threats including invasive species

Biological Weapons Convention prohibit development, production, http://www.fas.harvard.edu/~hsp/biologic.html (BWC) stockpiling, acquisition or retention of microbial or other biological agents that are not for peaceful use

Convention on International regulate invasive species not http://international.fws.gov/laws/citestxt.html Trade in Endangered Species covered by other agreements (CITES)

Convention on Migratory prevent, reduce or control factors http://www.cms.int/ Species of Wild Animals that endanger any species, including introduced exotic species

Convention on the Conservation prevent changes or minimize the http://sedac.ciesin.org/entri/texts/antarctic.marine.resources.1980.html of Antarctic Marine Living risk of changes in the Antarctic Resources marine ecosystem including the effects of alien species

United Nations Convention on prevent, reduce and control the http://www.univie.ac.at/RI/KONTERM/intlaw/konterm/vrkon_en/ the Law of the Sea (UNCLOS) intentional or accidental introduc- html/doku/unclos.htm tion of potentially harmful species to the marine environment

Protocol to the Antarctic Treaty require permits for introducing http://sedac.ciesin.org/entri/texts/antarctic.treaty.protocol.1991.html on Environmental Protection species to Antarctica

Convention on Biological prevent introductions or control or http://www.biodiv.org/ Diversity (CBD) eradicate alien or modified species that threaten ecosystems, habitats, species or human health

Framework Convention on stabilize and eventually reduce http://unfccc.int/ Climate Change greenhouse gas concentrations that raise temperatures, change rainfall patterns and induce or exacerbate invasions

Agreement on the Application of protect animals or plants from http://www.wto.org/english/tratop_e/sps_e/spsund_e.htm Sanitary and Phytosanitary pests and diseases Measures (SPS Agreement)

Agreement concerning Coopera- apply quarantine to prevent the http://sedac.ciesin.org/entri/texts/quarantine.of.plants.1959.html tion in the Quarantine of Plants introduction of plant pests, and their Protection against Pests diseases and weeds and Diseases

16 A PRIMER ON IINVASIVE SSPECIES IN CCOASTAL AND MMARINE WWATERS Agreement Relevance Web site

Convention on the Law of Non- prevent the introduction of species http://www.un.org/law/ilc/texts/nnavfra.htm navigational Uses of International into an international watercourse Watercourses

Biosafety Protocol (Protocol to control transboundary movement http://www.biodiv.org/biosafety/ the CBD) of living, modified organisms

European and Mediterranean prevent introduction and spread of http://www.eppo.org Plant Protection Organization pests and diseases of plants and animals

Protocol Concerning Specially prevent intentional or accidental http://www.cep.unep.org/pubs/legislation/spaw.html Protected Areas Wildlife to the introduction of harmful Convention for the Protection nonindigenous or genetically and Development of the Marine altered species into the Caribbean Environment of the Wider Region Caribbean Area

North American Agreement on develop recommendations http://www.cec.org Environmental Cooperation regarding harmful exotic species

North American Free Trade require notice of intent to restrict http://www.sice.oas.org/tradee.asp Agreement (NAFTA) imports

Convention on Great Lakes control and eradicate the non- http://www.glfc.org/pubs/conv.htm Fisheries Between the United native, highly invasive Atlantic sea States and Canada lamprey from the Great Lakes

United Nations Conference on control invasive species and their http://www.igc.org/habitat/agenda21/ Environment and Development impacts (UNCED)

World Trade Organization require notice of intent to restrict http://www.wto.org/wto/english/docs_e/legal_e/final_e.htm (WTO) – formerly GATT imports

International Maritime deal with ballast water as a vector http://www.imo.org organization for invasive species

United Nations Conference on protect forests against invasive http://www.un.org/documents/ga/conf151/aconf15126- Environment and Development species 3annex3.htm (UNCED)

Global Programme of Action for recognize invasive species as an http://www.unep.org/unep/gpa/pol2a.htm the Protection of the Marine important threat Environment from Land-based Activities

International Civil Aviation reduce risk of introductions http://www.icao.int/ Organization (ICAO) Resolution

A PRIMER ON INVASIVE SPECIES IN COASTAL AND MARINE WATERS 17 Appendix B. Experts on invasive species in the United States. (Source U.S. Fish and Wildlife Service; compiled November 2003).

ST Name and Title Agency Address Telephone Facsimile e-mail address

AL Vernon Minton Alabama Department 21055 Mildred Casey 334-968-7576 334-968-7307 [email protected] of Conservation & Gulf Shores, AL 36547 Natural Resources John Carlton Alabama Department 2204 Perimeter Road 334-450-3400 334-479-2593 [email protected] Chief, Field Operations of Environmental Mobile, AL 36615 Division Management

AK Bob Piorkowski Alaska Department of P.O. Box 25526 907-465-6109 907-465-2604 [email protected] Fish and Game Juneau, AK 99802 John Sisk (alternate) Office of the Governor P.O. Box 110001 907-465-3500 907-465-3532 [email protected] Juneau, AK 99811-0001

AZ Larry Riley Arizona Game and 2221 West Greenway Road 602-789-3258 602-789-3265 [email protected] Fish Commission Phoenix, AZ 85023-4399 Marc Dahlberg Arizona Game and 2221 West Greenway Road 602-789-3260 602-789-3265 [email protected] (alternate) Fish Commission Phoenix, AZ 85023-4399

AR Brian Wagner Arkansas Game and 915 East Sevier Street 877-847-2690 501-776-8362 [email protected] Fish Commission Benton, AR 72015 ext. 22

CA Susan Ellis California Department 1416 Ninth Street 916-653-8983 916-653-8256 [email protected] of Fish and Game Sacramento, CA 95814 Nate Dechoretz California 1220 N. Street 916-654-0768 916-653-2403 [email protected] (alternate) Department of Food Room A-357 and Agriculture Sacramento, CA 95814

CO Gary Skiba Colorado Division of 6060 Broadway 303-291-7466 303-291-7104 [email protected] Wildlife Denver, CO 80216

CT Arthur Rocque Connecticut Department 79 Elm Street 860-424-3001 860-424-4051 [email protected] Commissioner of Environmental Hartford, CT 06106 Protection

DE Jeff Tinsman Delaware Division of 89 Kings Highway 302-739-4782 302-739-6780 [email protected] Fish and Wildlife Dover, DE 19901

FL Lynn Griffin Florida Department of 3900 Commonwealth Blvd. 850-487-2231 850-922-5380 [email protected] Environmental Environmental MS 47 Administrator Protection Tallahassee, FL 32399-3000 Kurtis Gress (alternate) Florida Department of 3900 Commonwealth Blvd. 850-487-2231 850-922-5380 [email protected] Environmental Environmental MS 47 Administrator Protection Tallahassee, FL 32399-3000 Daniel Roberts Jr. Florida Fish and 100 Eighth Avenue SE 727-896-8626 603-971-2386 [email protected] Research Scientist Wildlife Conservation St. Petersburg, FL 33701 Commission

GA Ted Hendrickx Georgia Department 2123 U.S. Hwy. 278 SE 770-918-6418 706-557-3040 [email protected] of Natural Resources Social Circle, GA 30025

HI Domingo Cravalho Jr. Hawaii Department of 701 Ilalo Street, 808-586-0858 808-586-0864 [email protected] Agriculture, Plant Honolulu, HI 96813-5524 Quarantine Branch

ID Bill Horton Idaho Department of 600 S. Walnut, P.O. Box 25 208-334-3791 208-334-2114 [email protected] Fishery Coordinator, Fish and Game Boise, ID 83707 Anadromous Fish Fred Partridge Idaho Department of 600 S. Walnut, P.O. Box 25 208-334-3791 208-334-2114 [email protected] Fishery Coordinator, Fish and Game Boise, ID 83707 ext. 234 Resident Fish Glen Secrist Idaho State 2270 Old Penitentiary Rd. 208-332-8536 208-334-4062 [email protected] Chief, Bureau of Department of Boise, ID 83712 Vegetation Mgmt. Agriculture

18 A PRIMER ON IINVASIVE SSPECIES IN CCOASTAL AND MMARINE WWATERS ST Name and Title Agency Address Telephone Facsimile e-mail address

IL Mike Conlin Illinois Department of 600 North Grand Ave. 217-782-6424 217-785-8262 [email protected] Chief, Natural Resources West Springfield, IL 62706 Division of Fisheries Resources Pat Charlebois Illinois Natural History 400 17th Street 847-872-0140 847-872-8679 [email protected] (alternate) Survey, Lake Michigan Zion, IL 60099 Vice Chair, Biological Station Great Lakes Panel

IN Gwen White Indiana Department of 402 West Washington St. 317-232-4093 317-232-8150 [email protected] Program Staff Natural Resources, Division 273W 317-233-5468 Specialist of Fish and Wildlife Indianapolis, IN 46204 Randy Lang (alternate) Indiana Department of 402 West Washington St. 317-232-4094 317-232-8150 [email protected] Natural Resources, Division 273W of Fish and Wildlife Indianapolis, IN 46204

IA Marion Conover Iowa Department of Natural 502 9th Street 515-281-5208 515-281-6794 [email protected] Chief of Fisheries Resources Des Moines, IA 50319 Kim Bogenschutz Iowa Department of Natural 1436 255th Street 515-432-2823 515-432-2835 [email protected] (alternate) Resources Boone, IA 50036

KS Tom Mosher Kansas Department of P.O. Box 1525 316-342-0658 316-342-6248 [email protected] Wildlife and Parks Emporia, KS 66901

KY Ted Crowell Kentucky Department of 1 Game Farm Road 502-564-7109 502-564-6508 [email protected] Fish and Wildlife Resources Frankfort, KY 40601 ext. 467

LA Len Bahr Office of the Governor 625 North Fourth Street 225-342-3968 225-342-5214 [email protected] Executive Assistant Suite 1127 Baton Rouge, LA 70802 Dugan S. Sabins Environmental Program P.O. Box 82178 225-765-0246 225-765-0617 [email protected] (alternate) Louisiana Department of Baton Rouge, LA 70810 Environmental Quality 225-765-2336 225-765-5176 [email protected] Mark McElroy Louisiana Department of 2000 Quail Drive Wildlife and Fisheries Baton Rouge, LA 70898

ME Roy Bouchard Maine Department of State House Station 17 207-287-7798 207-287-7191 [email protected] Biologist III Environmental Protection Augusta, ME 04333 Peter Thayer Maine Department of P.O. Box 8 207-633-9539 207-633-9579 [email protected] Marine Resources Marine Resources Boothbay, ME 04575 Specialist John McPhedran Department of Environmental State House Station 17 207-287-6110 207-287-7191 [email protected] Invasive Species Protection, Bureau of Land Augusta, ME 04333-0017 Coordinator and Water Quality

MD Karen Eason Maryland Department of the 2500 Broening Highway 410-631-3611 410-631-3873 [email protected] Environment Baltimore, MD 21224 Edith Thompson Maryland Department of 580 Taylor Ave., E-1 410-260-8555 410-260-8596 [email protected] Invasive Species Natural Resources Annapolis, MD 21401 Policy Coordinator

MA Susan Snow-Cotter Massachusetts Coastal Zone 251 Causeway Street 617-626-1202 617-626-1240 susan.snow-cotter Assistant Director Management Program Suite 900 @state.ma.us Boston, MA 02114 Jim Straub Massachusetts Department 131 Barnum Road 508-792-7716 [email protected] of Environmental Building 3701 Management Devens, MA 01432 Jason Baker Executive Office of 251 Causeway Street 617-626-1204 617-626-1240 [email protected] Nonpoint Monitoring Environmental Affairs, Suite 900 Coordinator Office of Coastal Zone Boston, MA 02114-1219 Management

A PRIMER ON INVASIVE SPECIES IN COASTAL AND MARINE WATERS 19 ST Name and Title Agency Address Telephone Facsimile e-mail address

MI Roger Eberhardt Michigan Department of P.O. Box 30473 517-335-4227 517-335-4053 [email protected] Office of the Great Environmental Quality Lansing, MI 48909-7973 Lakes Emily Finnel (alternate) Michigan Department of P.O. Box 30473 517-335-7927 517-335-4053 [email protected] Office of the Great Environmental Quality Lansing, MI 48909-7973 Lakes

MN Jay Rendall Minnesota Department of 500 Lafayette Road 651-297-1464 651-297-7272 [email protected] Exotic Species Program Natural Resources, Fish and St. Paul, MN 55155-4020 Coordinator Wildlife Division Doug Jensen (alternate) Minnesota Sea Grant 2305 East 5th Street 218-726-8712 218-726-6556 [email protected] Exotic Species College Program Duluth, MN 55812-1445 Information Center Coordinator

MS Phil Bass Mississippi Department of P.O. Box 10385 601-961-5100 601-961-5337 [email protected] Director, Environmental Quality Jackson, MS 39204 Office of Pollution Control Gale Martin (alternate) Mississippi Soil and Water P.O. Box 23005 601-354-7645 601-354-6628 [email protected] Executive Director Conservation Commission Jackson, MS 39225-3005 David Felder Mississippi Department of 4279 Forest Park 601-664-3987 601-664-3938 [email protected] Biologist, Environmental Quality Jackson, MS 39211 Environmental Scientist II

MO Sue Bruenderman Missouri Department of 1110 College Avenue 573-882-9880 573-882-4517 [email protected]. Natural Resources Columbia, MO 65201 ext. 3239 state.mo.us

MT Tim Gallagher Montana Department of P.O. Box 200701 406-444-2448 406-444-4852 [email protected] Resource Program Fish, Wildlife and Parks Helena, MT 59620 Barbra Mullin Montana Department of P.O. Box 200201 406-444-3140 406-444-7336 [email protected] State Weed Agriculture, Agriculture Helena, MT 59620-0201 Coordinator Science Division

NE Steve Schainost Nebraska Game and Parks 2200 N. 33rd Street 402-471-5443 402-471-5528 [email protected] Commission Lincoln, NE 68503

NV Anita Cook Nevada Division of Wildlife 1100 Valley Road 775-688-1532 775-688-1595 [email protected] Reno, NV 89512 Jon Sjoberg Nevada Division of Wildlife 4747 Vegas Drive 702-486-5127 702-486-5133 Las Vegas, NV 89108

NH Carol Henderson New Hampshire Depart- 2 Hazer Drive 603-271-6649 603-271-6938 [email protected] Ecologist ment of Fish and Game Concord, NH 03301 Amy Smagula New Hampshire Depart- 6 Hazer Drive 603-271-2248 603-271-7894 [email protected] Clean Lakes and Exotic ment of Environmental P.O. Box 95 Species Program Services Concord, NH 03302 Coordinator

NJ Floyd Yoder New Jersey Department of P.O. Box 330 609-292-6075 609-292-4710 [email protected] Supervisor, Agriculture, Division of Trenton, NJ 08625 Seed Certification and Plant Industry Control Robert Cartica Office of Natural Lands 22 South Clinton Avenue 609-984-1015 609-984-1427 [email protected] Coordinator Management P.O. Box 404 Trenton, NJ 08625 David Snyder Office of Natural Lands 22 South Clinton Avenue 609-984-7849 609-984-1427 [email protected] Botanist Management P.O. Box 404 Trenton, NJ 08625

NM Brian Lang New Mexico Department of 1085-A Richards Avenue 505-476-8108 505-827-9956 [email protected] Game and Fish Santa Fe, NM 87505

20 A PRIMER ON IINVASIVE SSPECIES IN CCOASTAL AND MMARINE WWATERS ST Name and Title Agency Address Telephone Facsimile e-mail address

NY Timothy J. Sinnott New York Department of 625 Broadway 518-402-8970 518-402-9027 [email protected] Biologist 2 Environmental Albany, NY 12233-4756 Conservation Gerald A. Barnhart New York Department of 625 Broadway 518-457-5691 518-457-0341 [email protected] Assistant Director of Environmental Albany, NY 12233-4756 Fish and Wildlife Conservation William J. Culligan New York State Depart- 178 Point Drive North 716-366-0228 716-366-3743 [email protected] (alternate) ment of Environmental Dunkirk, NY 14048-1031 Supervising Aquatic Conservation, Lake Erie Biologist Fisheries Unit

NC Dean Horkavy North Carolina Division of 1611 Mail Service Center 919-715-5452 919-733-3558 [email protected] Water Resources Raleigh, NC 27699-1611

ND Terry Steinwand North Dakota Game and 100 N. Bismark Expressway 701-328-6313 701-328-6352 [email protected] Chief of Fisheries Fish Department Bismark, ND 58501-4095 Lynn Schlueter North Dakota Game and 7928 45th Street, NE 701-662-3617 701-662-3618 [email protected] (alternate) Fish Department Devils Lake, ND 58301

OH Gary Isbell Ohio Department of 1840 Belcher Drive, G 614-265-6345 614-262-1143 [email protected] Executive Admin., Natural Resources Columbus, OH 43224 Fish Management and Research Randy Sanders Ohio Department of 1840 Belcher Drive, G3 614-265-6344 614-262-1143 [email protected] (alternate) Natural Resources, Division Columbus, OH 43224 Aquatic Nuisance of Wildlife Species Program Administrator, Fish Management and Research

OK Jeff Boxrucker Oklahoma Department of 500 East Constellation 405-325-7288 [email protected] Wildlife Conservation Norman, OK 73072

OR Andrew Schaedel Oregon Department of 811 SW 6th Avenue 503-229-6121 503-229-6957 [email protected] Environmental Quality Portland, OR 97204 Mark Sytsma Portland State University, 1025 Harrison Street 503-725-3833 503-725-3888 [email protected] Aquatic Nuisance Center for Lakes and Portland, OR 97207-0751 Species Coordinator Reservoirs Tim Butler Oregon Department of 635 Capital Street, NE 503-986-4625 503-986-4737 [email protected] Manager, Agriculture Salem, OR 97301-2532 Noxious Weeds Control Program

PA Kelly Burch Pennsylvania Department of 230 Chestnut Street 814-332-6816 814-332-6125 [email protected] Chief, Environmental Protection Meadville, PA 16335 Office of the Great Lakes Eric Obert Penn State, Erie The Behrend College 814-898-6420 814-898-6462 ecol@psu-edu Program Director, Station Road Pennsylvania Sea Grant Erie, PA 16563-0101 Leo Dunn Pennsylvania Department of 2301 North Cameron St. 717-783-8462 717-787-5643 [email protected] Agriculture, Bureau of Room 311 Market Development Harrisburg, PA 17110

RI Kevin R. Cute Rhode Island Coastal 4808 Tower Hill Road 401-783-3370 401-783-3767 [email protected] Marine Resource Resource Management Wakefield, RI 02879 Specialist Council

SC Steve de Kozlowski South Carolina Department 207 Canterfield Road 803-734-9114 803-734-9200 [email protected] of Natural Resources, Land Columbia, SC 29212 and Water Conservation

A PRIMER ON INVASIVE SPECIES IN COASTAL AND MARINE WATERS 21 ST Name and Title Agency Address Telephone Facsimile e-mail address

SD Cliff Stone South Dakota Department 1125 N. Josephine Street 605-734-4538 605-734-6691 [email protected] of Game, Fish and Parks Chamberlain, SD 57325

TN William C. Reeves Tennessee Wildlife P.O. Box 40747 615-781-6575 615-781-6667 [email protected] Resources Agency Nashville, TN 37204

TX Bruce A. Moulton Texas Natural Resources P.O. Box 13087 512-239-4809 512-239-6195 [email protected] Program Specialist Conservation Commission Austin, TX 78711-3087 Larry McKinney Texas Parks and Wildlife 4200 Smith School Road 512-389-4636 512-389-4394 [email protected] (alternate) Department Austin, TX 78744 Earl Chilton Texas Parks and Wildlife 4200 Smith School Road 512-389-4652 512-389-4405 [email protected] Department Austin, TX 78744

UT Randy Randant Utah Division of Wildlife 1594 West North Temple 801-538-4760 801-538-4745 [email protected] Resources Salt Lake City, UT 84116

VT Michael Houser Vermont Department of 103 South Main Street 802-241-3777 802-241-3287 [email protected] Aquatic Biologist Environmental Protection Building 10N Waterbury, VT 05671-0408

VA Tom Wilcox Virginia Department of 4010 West Broad Street 804-367-8998 804-367-2427 [email protected] Game and Inland Fisheries Richmond, VA 23230

WA Scott Smith Washington Department of 600 Capital Way North 360-902-2724 360-902-3845 [email protected] Aquatic Nuisance Fish and Wildlife Olympia, WA 98501-1091 Species Coordinator Ron D. Shultz Washington Governor’s P.O. Box 43113 360-902-0676 360-902-0411 [email protected] (alternate) Office Olympia, WA 98501-3113 Executive Policy Assistant to the Governor Pam Meacham Washington Department of 600 Capital Way North 360-902-2741 360-902-3845 [email protected] Assistant Aquatic Fish and Wildlife Olympia, WA 98501-1091 Nuisance Species Coordinator Kathy Hamel Washington Department of P.O. Box 47600 360-407-6562 360-407-6426 [email protected] Aquatic Plant Specialist Ecology Olympia, WA 98504-7600

WV Brian McDonald West Virginia Division of P.O. Box 67 304-637-0245 304-637-0250 [email protected] Coordinator Natural Natural Resources Elkins, WV 26241 Heritage Program

WI Ron Martin Wisconsin Department of 101 S. Webster 608-266-9270 608-267-2800 [email protected] Chair, Natural Resources P.O. Box 7921 Great Lakes Panel Madison, WI 53707 Philip B. Moy Wisconsin Sea Grant 705 Viebahn Street 920-683-4697 920-683-4776 [email protected] (alternate) Advisory Service Manitowoc, WI 54220 Fisheries Specialist

WY Mike Stone Wyoming Game and Fish 5400 Bishop Boulevard 307-777-4559 307-777-4611 [email protected] Chief of Fisheries Department Cheyenne, WY 82006

DC Jon Siemien District of Columbia 51 N. Street, N.E. 202-535-2273 202-535-1373 [email protected] Department of Health, Room 5003 Fisheries and Wildlife Washington D.C. 20002 Division

22 A PRIMER ON IINVASIVE SSPECIES IN CCOASTAL AND MMARINE WWATERS Appendix C. Sources of information on introduced or invasive species, their management, and their use in research and education.

Description Source

Aquatic and marine species, all states http://nas.er.usgs.gov http://invasions.si.edu/ http://www.anstaskforce.gov/ http://www.sgnis.org/

Marine species, New England only http://massbay.mit.edu/exoticspecies

Aquatic species, Gulf of Mexico states http://nis.gsmfc.org/

Aquatic species, California only http://www.elkhornslough.org/invader.htm

Plants and insects, all states http://www.invasivespecies.org

Plants, all states http://invasives.eeb.uconn.edu

National Sea Grant College Program Invasive http://www.aquaticinvaders.org/nan_1d.cfm Species Clearinghouse

National Invasive Species Council http://www.invasivespecies.gov

National Invasive Species Council tool kit for http://www.invasivespecies.gov/toolkit/main.shtml managers

National Biological Information Infrastruc- http://www.nbii.gov/geographic/us/state.html ture federal and state resources

National Biological Information Infrastruc- http://www.nbii.gov/datainfo/bestpractices ture best management practices

Guidelines for working with microorganisms National Institutes of Health (NIH). 1968. Guidelines for Research Involving Recombinant DNA Molecules. Published in the Federal Register May 7, 1986 (51FR 16958-16961) with additional major actions published on August 24, 1987 (52F 31838); July 29, 1988 (53FR 28819); October 26, 1988 (53FR 43410); March 13, 1989 (54FR 10508); March 1, 1990 (55FR 7438); and August 11, 1987 (52FR 29800) with appendix P for plants and Q for animals. U.S. Department of Agriculture (USDA). 1984. Coordinated Framework for Regulation of Biotechnology. Published in the Federal Register December 31, 1984 (49FR 50856) and June 26, 1986 (51FR 23302+). U.S. Department of Agriculture (USDA). 1986. Advance Notice of Proposed USDA Guidelines for Biotechnology Research. Published in the Federal Register June 26, 1986 (51FR 23367-23393) and February 1, 1991 (56FR 4134-4149). U.S. Department of Agriculture (USDA). 1986. Introduction of Organisms and Products Altered or Produced Through Genetic Engineering Which are Plant Pests or for Which There is Reason to Believe are Plant Pests. Published in the Federal Register June 26, 1986 (51FR 23352-23366) and June 16, 1987 (52FR 22892-22915). Coulson, J. R., and R. S. Soper. 1989. Protocols for the Introduction of Biological Control Agents in the U.S. Chapter I, pages 2-35 In: Kahn, R. P. (ed.). Plant Protection and Quarantine. Volume III Special Topics. CRC Press, Inc., Boca Raton, Florida. U.S. Department of Agriculture (USDA). Office of Agricultural Biotechnology. 1988. USDA Guidelines for Research Outside the Laboratory Involving Biotechnology. Also Federal Register June 26, 1986 (51FR 23367-23313) and February 1, 1991 (56FR 4134-4149).

A PRIMER ON INVASIVE SPECIES IN COASTAL AND MARINE WATERS 23 Description Source

Guidelines for working with whole plants Jennings, D. P., and J. A. McCann. 1991. Recommended Protocol for Handling and animals Nonindigenous Aquatic Species. National Fisheries Research Center, U.S. Fish and Wildlife Service, Gainesville, FL. 43-page manuscript. http://www2.mcgill.ca/biology/phytotron/ipm_responsibilities.htm

International guidelines for working with European Inland Fisheries Advisory Commission. 1988. Code of Practice and Manual of nonindigenous species Procedures for Consideration of Introductions and Transfers of Marine and Freshwater Organisms. FAO. EIFAC. Occasional paper No. 23. 52 pages. International Council for the Exploration of the Sea. 1982. Proposed Guidelines for Implementing the ICES Code of Practice Concerning Introduction and Transfer of Marine Species. 23-page manuscript.

Guidelines for working with pathogens Anonymous. 1989. Operating Procedures for the Alma Quarantine Facility. Prepared for the Alma Research Station, Guelph, Ontario, Canada. 16 pages typewritten. Horner, R. W., and R. L. Eschenroder. 1991. Protocols to Minimize the Risk of Introducing Salmonid Disease Agents with Importation of Salmonid Fishes. Draft manuscript. 11 pages. Prepared for Great Lakes Fish Disease Control Committee. Pages 27-37. U.S. Department of Health and Human Services. 1984. Biosafety in Microbiological and Biomedical Laboratories. 1st Edition (March 1984). U.S. Department of Health and Human Services, Public Health Service, Centers for Disease Control, Atlanta, Georgia 30333, and National Institutes of Health, Bethesda, Maryland 20892. An additional 17 references on laboratory disease and pathogen control methods can be found listed in the Federal Register, May 7, 1986 (51FR 16965).

24 A PRIMER ON IINVASIVE SSPECIES IN CCOASTAL AND MMARINE WWATERS Educational Materials on Marine Invasives Preventing or controlling introductions is the best defense against invasive species. Florida Sea Grant and its partners provide these additional educational materials for coastal residents.

Can We Stop “Killer Algae” from Invading Florida? Several species of Caulerpa grow in Florida, and the state has ideal conditions for an introduction of the now infamous invasive alga, Caulerpa taxifolia, which decimated habitats in the Mediterranean. Florida’s coastal residents can help prevent an invasion of this killer algae by following a few simple measures outlined in this full-color, 2-page fact sheet. Contact Florida Sea Grant, (352) 392-2801, http://www.flseagrant.org. Single copies free, multiple copies have a charge. Request SGEF 155.

Are Native and Nonindigenous Seaweeds Overgrowing Florida’s East Coast Reefs? Scientists are concerned that extensive blooms of the non-native, green alga Caulerpa brachypus will spread on Florida’s reefs unless the amount of nutrients entering the sea can be reduced. This 2-page, color fact sheet describes macroalgal blooms that have been found on Florida’s east coast reefs and ways coastal residents can work to lower the nutrient loads that fuel them. Contact Florida Sea Grant, (352) 392-2801, http://www.flseagrant.org. Single copies free, multiple copies have a charge. Request SGEF 156.

Don’t Release Non-Native Species — Water Gardeners/Aquarium Hobbyists Most aquarium and water garden plants and animals sold in pet shops are non-natives imported predominantly from Central and South America, Africa, and Southeast Asia. This pair of two-sided, color rack cards explains the perils of releasing these non-native plants and animals into waterways and gives hobbyists safe alternatives for disposal. Contact Florida Sea Grant, (352) 392-2801, http://www.flseagrant.org. Single copies free, multiple copies have a charge.

Invasive Non-Native Plants Photo-Murals Two large, fully laminated photo-murals depict 75 invasive non-native aquatic and terrestrial plants found in Florida and throughout the U.S. All plants are depicted in large, strikingly attractive color photographs. Free to Florida teachers and public agency trainers who request them in writing, on letterhead from APIRS Photo-Mural, Center for Aquatic and Invasive Plants, 7922 NW 71 ST, Gainesville, FL 32653. Copies can be purchased from the IFAS Extension Bookstore, (800) 226-1764, http://www.ifasbooks.ufl.edu.

Divers Alert! Dive Card You can help prevent the spread of harmful marine invasives by reporting sightings of introduced species. This full-color, laminated card describes a variety of plants and animals that are already present in Florida waters or likely to invade soon. It also gives divers and boaters the Aquatic Nuisance Species toll free hotline and web site.

Green Mussel Alert Wallet Card Invasive green mussels appeared in Florida in 1999. These bright green shellfish can grow densely on boats, buoys, and piers, and they may become pests that crowd out native oysters. This wallet card illustrates mussels and gives instructions for reporting their presence. Contact Florida Sea Grant, (352) 392-2801, http://www.flseagrant.org. Single copies free, multiple copies have a charge.

3 Science Serving Florida’s Coast

PO Box 110409 University of Florida Gainesville, Florida 32611-0409

(352) 392-5870 www.FLSeaGrant.org

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