Summary Report of Freshwater Nonindigenous Aquatic Species in U.S

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Summary Report of Freshwater Nonindigenous Aquatic Species in U.S. Fish and Wildlife Service Region 4—An Update

April 2013

Prepared by: Pam L. Fuller, Amy J. Benson, and Matthew J. Cannister U.S. Geological Survey Southeast Ecological Science Center Gainesville, Florida

Prepared for: U.S. Fish and Wildlife Service Southeast Region Atlanta, Georgia

Cover Photos: Silver Carp, Hypophthalmichthys molitrix – Auburn University Giant Applesnail, Pomacea maculata – David Knott Straightedge Crayfish, Procambarus hayi – U.S. Forest Service

Table of Contents

Table of Contents ...... ii List of Figures ...... v List of Tables ...... vi INTRODUCTION ...... 1 Overview of Region 4 Introductions Since 2000 ...... 1 Format of Species Accounts ...... 2 Explanation of Maps ...... 3 References Cited ...... 4 CRUSTACEANS ...... 5 Overview ...... 5 Species Accounts of Crustaceans New to Region 4 ...... 11 Argulus japonicus Thiele, 1900 (Japanese Fishlouse) ...... 11 Elaphoidella bidens bidens (Schmeil, 1893) (a harpacticoid copepod) ...... 12 Mesocyclops ogunnus Onabamiro, 1957 (a cyclopoid copepod) ...... 13 Paracyclops bromeliacola Karaytug and Boxshall, 1998 (a copepod) ...... 14 Skistodiaptomus pallidus Herrick, 1879 (a calanoid copepod) ...... 15 Macrobrachium ohione (S. I. Smith, 1874) (Ohio Shrimp) ...... 16 Macrobrachium rosenbergii (De Man, 1879) (Giant River Prawn) ...... 17 Procambarus hayi (Faxon, 1884) (Straightedge Crayfish) ...... 19 Crustacean References Cited ...... 20 MOLLUSKS ...... 24 Overview ...... 24 Species Accounts of Mollusks New to Region 4 ...... 28 Dreissena bugensis Andrusov, 1897 (Quagga Mussel) ...... 28 Thiara scabra (Müller, 1774) (Pagoda Tiara) ...... 30 Viviparus georgianus (Lea, 1934) (Banded Mysterysnail)...... 31 Viviparus subpurpureus (Say, 1829) (Olive Mysterysnail) ...... 32 Mollusk References Cited ...... 33 AMPHIBIANS AND REPTILES ...... 35 Overview ...... 35

Species Accounts of Amphibians and Reptiles New to Region 4 ...... 43 Bombina orientalis (Boulenger, 1890) (Oriental Fire-bellied Toad) ...... 43 Afrixalus fornasini (Bianconi, 1849) (Fornasini’s Spiny Reed Frog) ...... 44 Kaloula pulchra Gray, 1831 (Malaysian Painted Frog or Chubby Frog) ...... 45 Pipa pipa (Linnaeus, 1758) (Suriname Toad) ...... 47 Amphiuma tridactylum Cuvier, 1827 (Three-toed Amphiuma) ...... 48 Cynops orientalis (Boie, 1826) (Oriental Fire-bellied Newt) ...... 49 Pachytriton labiatus (Unterstein, 1930) (Paddle-tail Newt) ...... 51 Paleosuchus palpebrosus (Cuvier, 1807) (Cuvier's Dwarf Caiman) ...... 52 Crocodylus acutus (Cuvier, 1807) (American Crocodile) ...... 53 Eunectes murinus (Linnaeus, 1758) (Green Anaconda) ...... 54 Eunectes notaeus (Cope, 1862) (Yellow Anaconda) ...... 55 Erpeton tentaculatus Lacepede, 1800 (Tentacled Snake)...... 57 Rhinoclemmys pulcherrima (Gray, 1855) (Painted Wood Turtle) ...... 58 Chelydra serpentina (Linnaeus, 1758) (Snapping Turtle) ...... 59 Macrochelys temminickii (Troost in Harlan, 1835) (Alligator Snapping Turtle) ...... 60 Glyptemys insculpta (LeConte, 1830) (Wood Turtle) ...... 61 Staurotypus salvinii Gray, 1864 (Pacific Coast Giant Musk Turtle) ...... 61 Apalone spinifera (Le Sueur, 1827) (Spiny Softshell) ...... 62 Amphibians and Reptiles References Cited ...... 64 FISH ...... 69 Overview ...... 69 Species Accounts of Fishes New to Region 4 ...... 84 Phenacogrammus interruptus (Boulenger, 1899) (Congo Tetra) ...... 84 Channa maculata (Lacepède, 1801) (Blotched Snakehead) ...... 85 Channa micropeltes (Cuvier in Cuvier and Valenciennes, 1831) (Giant Snakehead) ...... 86 Amphilophus labiatus (Günther, 1864) (Red Devil) ...... 86 Parachromis dovii (Günther, 1864) (Wolf Cichlid) ...... 87 Paraneetroplus melanurus x Paraneetroplus zonatus (hybrid cichlid) ...... 88 Paraneetroplus synspilus (Hubbs, 1935) (Redhead Cichlid) ...... 89 Mylopharyngodon piceus (Richardson, 1846) (Black Carp) ...... 90 Tanichthys albonubes (Lin, 1932) (White Cloud Mountain Minnow) ...... 93

iii

Apeltes quadracus (Mitchill, 1815) (Fourspine Stickleback) ...... 94 Gyrinocheilus aymonieri (Tirant, 1883) (Chinese Algae-eater) ...... 95 Atractosteus spatula (Lacepède, 1803) (Alligator Gar) ...... 95 Ancistrus sp. Kner, 1854 (Bristlenosed Catfish) possibly A. cf. cirrhosus ...... 96 Farlowella vittata Myers, 1942 (Stick catfish) ...... 98 Glyptoperichthys gibbiceps (Kner, 1854) (Leopard Pleco) ...... 99 Pterygoplichthys anisitsi Eigenmann and Kennedy, 1903 (Southern Sailfin Catfish) ...... 100 Osteoglossum bicirrhosum (Cuvier, 1829) (Arawana) ...... 101 Polypterus delhezi Boulenger, 1899 (Barred Bichir) ...... 102 Tetraodon nigroviridis (Marion de Procé, 1822) (Spotted Green Pufferfish) ...... 103 Fishes References Cited ...... 104

List of Figures

Figure 1. Number of new introduced species by taxonomic group...... 1 Figure 2. Number of new introduced species by continental origin...... 2 Figure 3. Number of new introduced species by vector...... 2 Figure 4. Total number of introduced crustacean species by taxonomic group...... 5 Figure 5. Total number of introduced crustacean species by continental origin...... 5 Figure 6. Total number of introduced crustacean species by vector...... 6 Figure 7. Total number of introduced crustacean species documented in 25-year intervals. (Note that the most recent interval is only 12 years.) ...... 6 Figure 8. Total number of introduced mollusk species by taxonomic group...... 24 Figure 9. Total number of introduced mollusk species by continental origin...... 24 Figure 10. Total number of introduced mollusk species by vector...... 25 Figure 11. Total number of mollusk species introduced in 25-year intervals. (Note that the current interval is only 12 years.) ...... 25 Figure 12. Total number of introduced A, amphibian andB, reptile species by taxonomic group...... 35 Figure 13. Number of new (since 2000) introduced A, amphibian and B, reptile (B) species by continental origin...... 35 Figure 14. Number of new (since 2000) introduced A, amphibian and B, reptile species by vector...... 36 Figure 15. Number of new fish species to the region by year...... 69 Figure 16. Number of new fish species to the region and state of first occurrence...... 69 Figure 17. Number of new introduced fish species by vector...... 70 Figure 18. Number of new introduced fish species by continental origin...... 70 Figure 19. Number of new state records per state 2001-2012...... 71 Figure 20. Number of new state records per year...... 71

List of Tables

Table 1. All nonindigenous crustacean species introductions into U.S. Fish and Wildlife Service Region 4...... 7 Table 2. All nonindigenous crustacean species introductions into U.S. Fish and Wildlife Service Region 4 by state...... 9 Table 3. All nonindigenous mollusk species introductions into U.S. Fish and Wildlife Service Region 4...... 26 Table 4. All nonindigenous mollusk species introductions into U.S. Fish and Wildlife Service Region 4 by state...... 27 Table 5. All nonindigenous amphibian and reptile introductions into U.S. Fish and Wildlife Service Region 4. . 37 Table 6. All nonindigenous amphibian and reptile introductions into U.S. Fish and Wildlife Service Region 4 by state...... 40 Table 7. New nonindigenous fish species introductions into U.S. Fish and Wildlife Service Region 4 since 2000...... 72 Table 8. All nonindigenous fish introductions into U.S. Fish and Wildlife Service Region 4 by state...... 73 Table 9. Fish species found as “new” in more than one state...... 83 Table 10. Sailfin catfish species sold as algae eaters to aquarists...... 83

INTRODUCTION

The southeast region of the United States including Caribbean territories (U.S. Fish and Wildlife Service (USFWS) Region 4) has a diverse fauna of fishes, amphibians, reptiles, mollusks, and crustaceans, and at the same time has some of the most imperiled fauna in the country (Williams and others, 1993; Taylor and others, 1996; Warren and others, 2000; Jelks and others, 2008). In addition to habitat degradation and loss, the region continues to have its waters inundated with exotic, or nonindigenous, species. Some of these may be of little consequence while others may cause substantial harm.

This report is an update through 2012 and builds upon the information in a similar report of the southeast region published in 2001 (Benson and others, 2001). The report is in the form of species profiles which contain information about identification, habitat, reproduction, distribution, impacts, and methods of introduction of each species introduced since 2000. This update is focused only on freshwater animals (fish, crustaceans, mollusks, amphibians, and reptiles), whereas the 2001 report included aquatic plants and marine animals.

A majority of the information for this report was derived from the U.S. Geological Survey’s Nonindigenous Aquatic Species Database program (http://nas.er.usgs.gov), which tracks the distribution of all non-native freshwater animals nationwide. For more information about the program and the sources used in compiling the database, please visit http://nas.er.usgs.gov/about/faq.asp. Overview of Region 4 Introductions Since 2000

There have been 49 new species introduced to Region 4 since 2000 (Figure 1). These are species that had never been collected from any state or territory in Region 4. One hundred and sixty species were recorded as new to a particular state since 2000. These are species that have been present somewhere in the southeast region, but were recorded in a state or territory where they had not been previously 4 documented—indicating either spread Fishes of an established population or new 7 19 Reptiles introductions. Two territories Crustaceans experienced the largest proportional increases relative to the states in the 8 Amphibians region: Puerto Rico (69 percent) and Mollusks Virgin Islands (60 percent). In terms of 11 total numbers of species, the two highest were Florida (47 percent) and Puerto Rico (36 percent). Fishes Figure 1. Number of new introduced species by taxonomic group. represent the largest taxonomic group.

Nearly equal numbers of these species are native to South and Central America, Asia and Eurasia, or other areas of North America whereas a much smaller fraction originates from Africa, and none from Australia (Fig. 2).

3 South and Central America 17 13 Asia/Eurasia

North America

Africa 16

Figure 2. Number of new introduced species by continental origin.

The two dominant vectors for species introductions are pet releases and aquarium/water garden releases. If these two categories are combined into a single category as “pet trade,” this accounts for 35 species, more than one-half of all the new species (Figure 3).

1 1 Aquarium/ water garden release 1 1 2 Pet release 4 Escaped aquaculture 18 Hitchhiker 4 Unknown Shipping Bait release Escaped farm 17 Food release

Figure 3. Number of new introduced species by vector.

Format of Species Accounts

The accounts are organized by taxonomic group. Within each group, they are further organized alphabetically by family, genus, and species. The report focuses on the group(s) or subgroup(s) that have been introduced the most and identifies the vector and source region of these species to more readily identify possible management actions. Accounts are given for each new species found in the region and

2 include the common and scientific name, a description of the native range and introduced range within Region 4, distribution maps, and information on vectors and impacts. We decided to capitalize all common names for consistency throughout the report, following convention used for fishes set forth by the American Fisheries Society (Nelson and others, 2013) and the convention for amphibians and reptiles as set forth by the Society for the Study of Amphibians and Reptiles (Moriarty, 2012). The names of mollusks and crustaceans used in this report follow those set forth in Turgeon and others (1998) and McLaughlin and others (2005), respectively. In the updated version of the Names of Fishes, common names are now capitalized. All scientific names are followed by the taxonomic authority who named them. When these names appear in parentheses it is because the genus has changed since it was originally described. Authorities without parentheses indicate the genus is the same as the original name.

For some species accounts, we used the common Web sites of AmphibianWeb, and Encyclopedia of Life because of the lack of accessible primary literature in English for these species.

It was determined that introductions of several species had been overlooked in the 2001 report; therefore, we included these omissions in this update. Explanation of Maps

Distributions are mapped using river drainages to relate locality data within natural drainage systems. River drainages are described by the U.S. Geological Survey’s (USGS) Hydrologic Unit Codes (HUC), a nested classification that identifies hydrologic regions at many levels in the United States. When possible, maps herein employ the 8-digit level HUC, where the species has been naturalized, collected, or observed. Each account has a U.S. map that depicts its introduced range (dark brown shaded 8-digit hydrologic units) across the entire country. Native ranges, when available, can be found on the maps, they are shaded in light brown. There also are regional maps that allow a more detailed look at the introduced ranges within Region 4 states. Large red circles aid the reader in finding very small hydrologic units. Small symbols on the region maps represent locations and its shape distinguishes its spatial accuracy: a circle is accurate, a square is approximate, and a triangle is a centroid. The color of these symbols represents the status of the introduction at that location: red is established, blue is failed, and green is unknown. States that are shaded entirely in pink represent an introduction in the state, but the location was not specified in the source documentation. The distribution information in this report was compiled from scientific literature, reports, personal communications, and natural history museum databases.

References Cited

Benson, A.J., Fuller, P.L., and Jacono, C.C., 2001, Summary report of nonindigenous aquatic species in U.S. Fish and Wildlife Service Region 4. U.S. Geological Survey, Gainesville, FL. Also available at: http://fl.biology.usgs.gov/Region_4_Report/index.html.

Jelks, H.L., Walsh, S.J., Burkhead, N.M., Contreras-Balderas, S., Diaz-Pardo, E., Hendrickson, D.A., Lyons, J., Mandrak, N.E., McCormick, F., Nelson, J.S., Platania, S.P., Porter, B.A., Renaud, C.B., Schmitter-Soto, J.J., Taylor, E.B., and Warren, M.L., 2008, Conservation status of imperiled North American freshwater and diadromous fishes: Fisheries, v. 33, no. 8, p. 372–407.

McLaughlin, P.L., and 39 others, 2005, Common and scientific names of aquatic invertebrates from the United States and Canada—Crustaceans: Bethesda, Maryland, American Fisheries Society Special Publication 31, 545 p.

Moriarty, J.J., ed., 2012, Scientific and standard English names of amphibians and reptiles of North America north of Mexico, with comments regarding confidence in our understanding (7th ed.): Society for the Study of Amphibians and Reptiles, 101 p. (Also available at http://home.gwu.edu/~rpyron/publications/Crother_et_al_2012.pdf.)

Nelson, J.S., Crossman, E.J., Espinosa-Perez, H., Findley, L.T., Gilbert, C.R., Lea, R.N., and Williams, J.D., 2004, Common and scientific names of fishes from the United States, Canada and Mexico (6th ed.): Bethesda, Maryland, American Fisheries Society, 386 p.

Taylor, C.A.; Warren, M.L., Jr.; Fitzpatrick, J.F., Jr.; Hobbs, H.H., III; Jezerinac, R.F.; Pflieger, W.L.; and Robison, H.W., 1996, Conservation status of crayfishes of the United States and Canada: Fisheries, v. 21, no. 4, p. 25–38.

Turgeon, D.D.; Quinn, J.F., Jr.; Bogan, A.E.; Coan, E.V.; Hochberg, F.G.; Lyons, W.G.; Mikkelsen, P.M.; Neves, R.J.; Roper, C.F.E.; Rosenberg, G.; Roth, Barry; Scheltema, A.; Thompson, F.G.; Vecchione, M.; and Williams, J.D., 1998, Common and scientific names of aquatic invertebrates from the United States and Canada—Mollusks (2nd ed.): Bethesda, Maryland, American Fisheries Society Special Publication 26, 526 p.

Warren, M.L., Jr.; Burr, B.M.; Walsh, S.J.; Bart, H.L., Jr.; Cashner, R.C.; Etnier, D.A.; Freeman, B.J.; Kuhajda, B.R.; Mayden, R.L.; Robison, H.W.; Ross, S.T.; and Starnes, W.C., 2000, Diversity, distribution, and conservation status of the native freshwater fishes of the southern United States: Fisheries, v. 25, no. 10, p. 7–31.

Williams, J.D.; Warren, M.L., Jr.; Cummings, K.S.; Harris, J.L.; and Neves, R.J., 1993, Conservation status of freshwater mussels of the United States and Canada: Fisheries, v. 18, no. 9, p. 6–22.

CRUSTACEANS

Overview

Overall, 32 crustacean species have been documented outside their historic range in U.S. Fish and Wildlife Service Region 4 dating back one century, eight of which were introduced into the southeastern United States region since 2000 (Table 1). The largest group of newly introduced species during the past decade is crayfishes, followed by copepods (Figure 4). Since 2000, most of the new 1 introductions are copepods, followed by 3 Crayfish freshwater shrimp, mysids, and cladocerans. 4 Copepods Although new species continue to be 14 introduced, only a few species were found to Shrimp have spread from states from which they were originally introduced (Table 2). Cladocerans 10

Mysids The majority of newly introduced species originate from North America (Figure 5); Figure 4. Total number of introduced crustacean however, two copepod species, one each species by taxonomic group. from Asia and South America, are new to USFWS Region 4 as well as to North America (Table 1).

2 1

3 North America

Asia/Eurasia

South and Central 18 8 America Australia

Africa

Figure 5. Total number of introduced crustacean species by continental origin.

Of greatest concern may be the crayfish because they commonly are distributed as bait, and introduced populations are known to outcompete natives ones. Bait 3 bucket releases and hitchhikers are the Bait release 3 most common pathway for the most 10 Hitchhiker recent introductions into the Region

(Figure 6). 6 Unknown

Stocked 10 Escaped aquaculture

Figure 6. Total number of introduced crustacean species by vector.

The number of new species documented since 2000 has the potential to surpass the number introduced in the previous 25 years (Figure 7). This increase most likely is from a combination of two factors: new introductions and better detection.

Number of New Species 18 16 14

12 10 8

6 4

2 0 1900-1925 1926-1950 1951-1975 1976-2000 2001-2012

Figure 7. Total number of introduced crustacean species documented in 25- year intervals. (Note that the most recent interval is only 12 years.)

Table 1. All nonindigenous crustacean species introductions into U.S. Fish and Wildlife Service Region 4. First Population Group Family Scientific Name Common Name Year Continent of Origin Likely Vector Status Cladocerans Daphnidae Daphnia lumholtzi water flea 1990 Australia/Asia hitchhiker with stocked fish Established Cladocerans Sididae Diaphanosoma brevireme † cladoceran 1960 South and Central America unknown Unknown Cladocerans Sididae Diaphanosoma fluviatile † cladoceran 1960 South and Central America unknown Unknown Copepods Argulidae Argulus japonicus Japanese Fishlouse 2003 Asia hitchhiker with stocked fish Established Copepods Canthocamptidae Elaphoidella bidens bidens copepod 2005 Eurasia hitchhiker with ballast water Unknown Copepods Cyclopidae Bryocyclops muscicola † copepod 1999 Asia hitchhiker on imported plants Unknown Copepods Cyclopidae Mesocyclops ogunnus * copepod 2007 Asia hitchhiker on imported plants Unknown Copepods Cyclopidae Mesocyclops pehpeiensis † copepod 1987 Asia hitchhiker on imported plants Unknown Copepods Cyclopidae Paracyclops bromeliacola * copepod 2005 South America hitchhiker on imported plants Unknown Copepods Diaptomidae Skistodiaptomus pallidus calanoid copepod 2005 North America hitchhiker with stocked fish Unknown Copepods Ergasilidae Neoergasilus japonicus † parasitic copepod 1993 Asia hitchhiker with stocked fish Unknown Copepods Lernaeidae Lernaea cyprinacea † anchor worm 1977 Asia unknown Unknown Copepods Temoridae Eurytemora affinis † calanoid copepod 1912 North America hitchhiker with ballast water Established Crayfish Cambaridae Cambarellus shufeldtii Cajun Dwarf Crayfish 1937 North America stocked Unknown Crayfish Cambaridae Cambarus cumberlandensis Cumberland Crayfish 1989 North America released bait Unknown Crayfish Cambaridae Cambarus longirostris Longnose Crayfish 1989 North America released bait Unknown Crayfish Cambaridae Faxonella clypeata Ditch Fencing Crayfish 1937 North America stocked Unknown Crayfish Cambaridae Orconectes neglectus chaenodactylus † crayfish 1998 North America released bait Established crayfish Cambaridae Orconectes palmeri creolanus Creole Painted Crayfish 1999 North America released bait Established Crayfish Cambaridae Orconectes placidus Placid Crayfish 1976 North America released bait Unknown Crayfish Cambaridae Orconectes rusticus † Rusty Crayfish 1989 North America released bait Unknown Crayfish Cambaridae Orconectes virilis Virile Crayfish 1980 North America released bait Unknown Crayfish Cambaridae Procambarus acutus acutus White River Crawfish 1938 North America released bait Unknown Crayfish Cambaridae Procambarus clarkii Red Swamp Crayfish 1954 North America released bait Unknown Crayfish Cambaridae Procambarus hayi Straightedge Crayfish 2009 North and Central America unknown Unknown Crayfish Cambaridae Procambarus seminolae Seminole Crayfish 1981 North America released bait Unknown Crayfish Parastacidae Cherax quadricarinatus Australian Redclaw 1998 Australia escaped captivity aquaculture Established

Table 1. All nonindigenous crustacean species introductions into U.S. Fish and Wildlife Service Region 4. (Continued) First Population Group Family Scientific Name Common Name Year Continent of Origin Likely Vector Status Mysids Mysidae Taphromysis louisianae† Mysid Shrimp 1981 North America unknown Unknown Shrimp Palaemonidae Macrobrachium macrobrachion Brackish River Prawn 1995 Africa escaped captivity aquaculture Unknown Shrimp Palaemonidae Macrobrachium ohione Ohio Shrimp 2005 North America unknown Unknown Shrimp Palaemonidae Macrobrachium olfersii † Bristled River Shrimp 1933 North America stocked Unknown Shrimp Palaemonidae Macrobrachium rosenbergii Giant River Prawn 2001 Asia escaped captivity aquaculture Unknown

Bold = New to U.S. Fish and Wildlife Service Region 4 since 2000 * = New to United States † = Introduced to USFWS Region 4 prior to 2001 but not included in last report

Table 2. All nonindigenous crustacean species introductions into U.S. Fish and Wildlife Service Region 4 by state/territory. Group Family Scientific Name Common Name AL AR FL GA KY LA MS NC PR SC TN VI Cladocerans Daphnidae Daphnia lumholtzi Water Flea X X X X X X X X X Cladocerans Sididae Diaphanosoma brevireme † cladoceran X Cladocerans Sididae Diaphanosoma fluviatile † cladoceran X X Copepods Argulidae Argulus japonicus Japanese Fishlouse N N N Copepods Canthocamptidae Elaphoidella bidens bidens copepod N N N N N N Copepods Cyclopidae Bryocyclops muscicola † copepod X Copepods Cyclopidae Mesocyclops ogunnus * copepod N Copepods Cyclopidae Mesocyclops pehpeiensis † copepod X X Copepods Cyclopidae Paracyclops bromeliacola * copepod N Copepods Diaptomidae Skistodiaptomus pallidus calanoid copepod N N N Copepods Ergasilidae Neoergasilus japonicus † parasitic copepod X Copepods Lernaeidae Lernaea cyprinacea † anchor worm X N Copepods Temoridae Eurytemora affinis † calanoid copepod X X X X X X Crayfish Cambaridae Cambarellus shufeldtii Cajun Dwarf Crayfish X Crayfish Cambaridae Cambarus cumberlandensis Cumberland Crayfish X Crayfish Cambaridae Cambarus longirostris Longnose Crayfish X Crayfish Cambaridae Faxonella clypeata Ditch Fencing Crayfish X Crayfish Cambaridae Orconectes neglectus chaenodactylus † crayfish X Crayfish Canbaridae Orconectes palmeri creolanus† Creole Painted Crayfish X Crayfish Cambaridae Orconectes placidus Placid Crayfish N X Crayfish Cambaridae Orconectes rusticus † Rusty Crayfish X X Crayfish Cambaridae Orconectes virilis Virile Crayfish X X X X Crayfish Cambaridae Procambarus acutus acutus White River Crawfish X X Crayfish Cambaridae Procambarus clarkii Red Swamp Crayfish X X X X Crayfish Cambaridae Procambarus hayi Straightedge Crayfish N Crayfish Cambaridae Procambarus seminolae Seminole Crayfish X Crayfish Parastacidae Cherax quadricarinatus Australian Redclaw X Mysids Mysidae Taphromysis louisianae † Mysid Shrimp X

Table 2. All nonindigenous crustacean species introductions into U.S. Fish and Wildlife Service Region 4 by state/territory. (continued) Group Family Scientific Name Common Name AL AR FL GA KY LA MS NC PR SC TN VI Shrimp Palaemonidae Macrobrachium macrobrachion Brackish River Prawn X Shrimp Palaemonidae Macrobrachium ohione Ohio Shrimp N Shrimp Palaemonidae Macrobrachium olfersii † Bristled River Shrimp X N X Shrimp Palaemonidae Macrobrachium rosenbergii Giant River Prawn N N Bold = New to U.S. Fish and Wildlife Service Region 4 since 2000 * = New to United States † = Introduced to USFWS Region 4 prior to 2001 but not included in last report X = Present in or prior to 2001 N = New since 2000

Species Accounts of Crustaceans New to Region 4

Argulus japonicus Thiele, 1900 (Japanese Fishlouse)

Life History: This small, brown ectoparasitic copepod ranges from 4 to 9 millimeters (mm) long and 3 to 6 mm wide (Pilgrim, 1967). Reproduction in dioecious Argulus japonicus is sexual and output is high. Although mating occurs on the body of the host fish, females lay 1–9 strings of 5–226 eggs in 1–6 rows on hard substrate, which they cover in gelatinous excretions (Shafir and Van As, 1986). Eggs hatch U.S. Geological Survey optimally at 20–30 degrees Celcius (°C ) in 2 to 4 weeks; eggs did not develop at temperatures of 9–11.5 °C (Shafir and Van As, 1986). Newly hatched larvae survive for 1 or 2 days on nutrition from their yolk sacs but soon require nutrition from the host’s epithelial cells and mucus (Tam and Avenant-Oldewage, 2006). After hatching, the rest of the life cycle (excluding egg-laying) takes place on the body of the host fish. Argulus japonicus lays eggs continuously; however, in winter, eggs may stay dormant until warmer temperatures conducive to hatching occur in spring (Shafir and Oldewage, 1992). As this species develops, it passes through approximately six different larval instar stages before metamorphosing into an adult. Adults can survive without a host for approximately 2 weeks (Avenant-Oldewage and Swanepoel, 1993). Being highly mobile, they leave the host fish regularly to breed or switch host fish; the lifespan is estimated to be 1 year (Shafir and Oldewage, 1992). The fishlouse feeds by inserting its stylet under the scale and drawing blood from the fish after piercing through the skin (Shafir and Oldewage, 1992).

Native Range: Argulus japonicus is native to Asia, where its common hosts, Goldfish (Carassius auratus) and Common Carp (Cyprinus carpio), also are native species (Mills and others, 1993; Bunkley-Williams and Williams, 1994).

Nonindigenous Range: Argulus japonicus is listed as being present in unspecified locations in Florida, Georgia, and Louisiana in the southeast United States as well as in California, Hawaii, Illinois, Maryland, Michigan, Washington, and Wisconsin (McLaughlin and others, 2005) (See pink shaded states in map below).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown

Vector: Argulus japonicus was very likely introduced to North America with its host Carassius auratus through the aquarium industry (Mills and others, 1993).

Impacts: Argulus japonicus can cause severe damage to the integument of its hosts, sometimes resulting in death by affecting the appetite of a fish, and subsequently affecting growth rates. This parasite attaches to the skin, fins, or gills by way of suckers, and then feeds. It also may aid in the transmission of Rhabdovirus carpio, the spring viraemia rhabdovirus; larval nematodes; and the fungal disease caused by Saprolegnia in some parts of the world (Gresty and others, 1993; Avenant-Oldewage, 2001). Introduced in a South African reservoir, this parasite infests a wide range of fishes (Kruger and others, 1983).

Comments: Gizzard Shad (Dorosoma cepedianum) is a host fish (Poly, 1998) as well as Yellow Perch (Perca flavescens), Koi Carp (Cyprinus carpio), and Goldfish (Carassius auratus) (Hudson and Lesko, 2003).

Elaphoidella bidens bidens (Schmeil, 1893) (a harpacticoid copepod)

Habitat: This species inhabits freshwater lakes, streams, reservoirs, and marshes; it can also tolerate eutrophic waters (Hudson and Lesko, 2003).

Life History: Copepods are small animals, less than 1 mm in length. Males are rarely encountered, and the most common mode of reproduction is parthenogenesis U.S. Geological Survey (Hudson and Lesko, 2003). This characteristic makes it different from all of its other North American congeners (Bruno and others, 2000).

Native Range: The genus has a worldwide distribution; however, Bruno and others (2000) believe it may have been introduced to North America from Eurasia because a large majority of records come from disturbed habitats.

Nonindigenous Range: Nonspecific locations in Florida, Georgia, Louisiana, North Carolina, South Carolina, and Tennessee (McLaughlin and others, 2005) (See pink shaded states in map below); also found in modest numbers in the lower Great Lakes (Huron, Erie, and Ontario) (Hudson and Lesko, 2003).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown 12

Pathway: Unknown, but because it has also been found in the Great Lakes, ballast water may have been how it was introduced initially.

Impacts: Unknown.

Comments: Adults are capable of surviving complete desiccation for months; the nauplii of this species have been found in mud taken from the feet of shore birds (Hudson and Lesko, 2003).

Mesocyclops ogunnus Onabamiro, 1957 (a cyclopoid copepod)

Habitat: This species is found in freshwater to oligohaline lakes and reservoirs (Hribar and Reid, 2008).

Life History: At 30 °C, this copepod develops from egg to adult in 8 days (Bonou and others, 1991 as cited in Hribar and Reid, 2008). They migrate vertically upward at night and back down during the day (Hribar and Reid, 2008). Mesocyclops sp. U.S. Geological Survey They feed on other copepods, cladocerans, rotifiers, and algae (Hribar and Reid, 2008). Younger copepodids primarily are herbivorous (Gophen, 1977 as cited in Hribar and Reid, 2008). Females can grow to 1.2 mm in size (Matsumura-Tundisi and Silva, 2002).

Native Range: Widespread in Africa and Asia (Hribar and Reid, 2008).

Nonindigenous Range: In 2007, this species was found in an artificial container and in a semi-permanent rain pond on Vaca Key, Florida (Hribar and Reid, 2008). (See green marker on map below)

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: The origins are unknown but it is highly likely it came to the United States through the ornamental plant trade. Some copepods have been known to be found in the leaves of bromeliads and in the soils of ornamental plants (Hribar and Reid, 2008). In South America, where it is also considered introduced, Matsumura-Tundisi and Silva (2002) believe it may have come with shipments of African fishes.

Impacts: Unknown. They are known to prey on mosquito larvae of the yellow fever mosquito, Aedes aegypti (L.), (Marten and Reid, 2007 as cited in Hribar and Reid, 2008). Mesocyclops ogunnus may be able to successfully compete with the native copepod, Mesocyclops edax (Hribar and Reid, 2008).

Comments: This is the first documentation of this species in the United States and was found during routine sampling for mosquito larvae. This copepod species can survive and reproduce in waters up to 5 parts per thousand (ppt) salinity (Bonou and others, 1991 as cited in Hribar and Reid, 2008).

Paracyclops bromeliacola Karaytug and Boxshall, 1998 (a copepod)

Habitat: This copepod is found in freshwater to oligohaline lakes, streams, and wetlands (Hribar and Reid, 2008).

Life History: Members of this genus are small, usually less than 1 mm in length. Reproduction is sexual with 6 nauplius and 5 copepodid stages to adulthood, females Paracyclops sp. U.S. Geological Survey can carry several egg sacs containing 8–36 eggs total, generation time is about 3–4 weeks, and continuous reproduction is year round (Karaytug and Boxshall, 1998).

Native Range: Brazil, South America (Karaytug and Boxshall, 1998).

Nonindigenous Range: Several dozen specimens were collected from the leaf cup of a bromeliad plant in 2005 at Key Colony Beach on Shelter Key in Florida (Reid and Hribar, 2006).

Impacts: Unknown.

Comments: Paracyclops species have been experimentally injected with microsporidian parasites of mosquitos as a possible biocontrol (Micieli and others, 2000). The leaf cups of bromeliads can provide an unusual microhabitat (Karaytug and Boxshall, 1998).

Skistodiaptomus pallidus Herrick, 1879 (a calanoid copepod)

Habitat: This species is found in freshwater ponds, lakes, reservoirs, and rivers (Torke, 2001).

Life History: Females are approximately 0.8–1.2 mm long, whereas males are about 0.7–1 mm long (Geddes and Cole, 1981; Torke, 2001; Hudson and Lesko, 2003). Breeding in general takes place from about March to November in its northern limit, U.S. Geological Survey where females can produce as many as 20 eggs per brood (Chapman and others, 1985; Hudson and Lesko, 2003). Skistodiaptomus pallidus produces some eggs that go through a diapause stage before they hatch, which may help mitigate the effects of a poor year in reproduction and recruitment (Wonham and others, 2005). Geiling and Campbell (1972) found it takes about 66 days at 10 °C and 15 days at 25 °C for development from the egg to the adult stage. Skistodiaptomus pallidus feeds on phytoplankton, especially individual algae greater than 53 microns (μm) in size (Geiling and Campbell, 1972) and can also selectively and intensely prey on some rotifer species (Geiling and Campbell, 1972; Williamson and Butler, 1986).

Native Range: Skistodiaptomus pallidus is native to the north-central plain states, southern United States in the Mississippi River Basin, Texas, and west to Colorado (Pennak, 1989; Mills and others, 1993; Torke, 2001). The home range is not depicted on the map below.

Nonindigenous Range: Unspecified locations in Georgia, North Carolina, and South Carolina (McLaughlin and others, 2005) (See pink shaded states on map below.).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown

Vector: Skistodiaptomus pallidus could have been introduced accidentally in bait buckets, on fishing equipment, by recreational boaters, with hatchery stockings from the Mississippi River Basin, or through dispersal (Mills and others, 1993).

Impacts: Skistodiaptomus pallidus is an efficient omnivorous predator, with the ability to prey on preferred rotifers and microzooplankton from large distances. It also consumes algae and practices cannibalism, which may allow populations to persist when resource availability is low (Williamson and Butler, 1986; Williamson and Vanderploeg, 1988). It also has been known to attain very high densities in suitable habitats, reaching 10,000 per cubic meter in a Lake Erie marsh to unknown consequences (Krieger and Klarer, 1991). Skistodiaptomus pallidus became the primary calanoid copepod in a particularly eutrophic part of Lake Tahoe, dominating two previously common species, Leptodiaptomus tyrrelli and Epischura nevadensis (Byron and Saunders, 1981). Additionally, based on evidence from an Ohio lake, it has been suggested that S. pallidus is an intermediate host for the parasitic worm Tanaorhamphus longirostris, although study of this occurrence has been limited (Hubschman, 1983). Documented evidence combined with its record of spread across the United States (Byron and Saunders, 1981) have led some recently colonized areas, like New Zealand, to express concern over the potential effects of S. pallidus on native ecosystems (Duggan and others, 2006).

Macrobrachium ohione (S. I. Smith, 1874) (Ohio Shrimp)

Habitat: This shrimp inhabits primarily freshwater rivers and streams; however, their life cycle is associated with estuarine waters for reproduction (Bowles and others, 2000).

Life History: This species is amphidromous, where adults migrate downstream to spawn near estuaries, and juveniles swim back upstream and live as adults in freshwater (Bowles and others, 2000). Females, which Ray Bauer are larger than males, can reach 110 mm in total length and are gray to green in color with light blue spots (Bowles and others, 2000). Females can produce 6,000–24,000 eggs with a lifespan of only 2 years (Huner, 1977).

Native Range: Macrobrachium ohione is the most widely distributed and abundant of the six river shrimp species in the United States (Bowles and others, 2000) and originally was described from the Ohio River. It also is found in the middle and lower Mississippi River from Illinois to Louisiana, west to Oklahoma and Texas, in the Atlantic drainage from Virginia to northern Florida, and in the Gulf region in Alabama and Mississippi (Bowles and others, 2000; Hobbs and Lodge, 2010).

Nonindigenous Range: Three individuals were collected in a sample in 1 day from Caloosahatchee River near Fort Myers, Florida in 2005.

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: Most likely the result of a bait bucket introduction. There has been a commercial fishery throughout its native range for bait as well as for human consumption for many years (Bowles and others, 2000; Bauer and Delahoussaye, 2008).

Impacts: Probably minimal impacts as this species is on the decline as a result of human impacts such as impoundments, river channelization, pollution, commercial fishing, and exotic predators (Barko and Hrabik, 2004; Bowles and others, 2000; Bauer and Delahoussaye, 2008; Hobbs and Lodge, 2010).

Comments: It is the only endemic species of the genus in North America (Hobbs and Lodge, 2010).

Macrobrachium rosenbergii (De Man, 1879) (Giant River Prawn)

Habitat: The Giant River Prawn is found in tropical freshwater lakes and rivers; however, estuaries play a role in reproduction in the native ranges of most Macrobrachium species [Food and Agriculture Organization of the United Nations (FAO), 2005–2012].

Life History: Males can grow to 320 mm and easily are recognized by their extremely long claws; females are Jarek Tuszynski smaller, reaching 250 mm total length (TL) (FAO, 2005– 2012). It is the largest of the genus (Mather and de Bruyn, 2003). Males deposit a packet of sperm on the female where they are then fertilized. After fertilization, gravid females migrate downstream to the estuary where the eggs hatch as free-swimming larvae (FAO, 2005–2012). Females can lay 10,000 to 50,000 eggs during each spawn, which happens several times a year (Wynne, 2000). The newly hatched larvae pass through 11 molts in 35 days in the estuary to become postlarval (less than one-half inch in length), then migrate back upstream to become adults (Wynne, 2000). The prawns are capable of walking on the substrate, on land at water edges, and up vertical surfaces such as small waterfalls (FAO, 2005–2012). They are omnivorous—feeding on plants, algae, mollusks, worms, and fish.

Native Range: Southern Asia from Pakistan to Vietnam, and east to northern Australia and Papua New Guinea (Mather and de Bruyn, 2003).

Nonindigenous Range: In slightly saline conditions, several dozen, including adult males, adult females, and juveniles, were collected from Simmons Bayou on the Gulf Coast of Mississippi in 2001 (Woodley and others, 2002). In Puerto Rico and the Caribbean region, there have been many mass releases of this widely cultured prawn where they are considered ecologically harmless (Williams and others, 2001).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown

Vector: Most likely these are escapes from aquaculture operations because it is widely cultured around the world (Mather and de Bruyn, 2003). It is not known whether the juveniles are escapes, the result of reproduction, or both (Woodley and others, 2002).

Impacts: In laboratory testing, this freshwater prawn was determined to be a carrier of, and tolerant to, the white spot syndrome virus that is known to affect marine shrimps (Hameed and others, 2000). White spot syndrome virus can cause high mortality in aquaculture operations. A disease outbreak in the United States could potentially affect economically and ecologically important native species of shrimp (Woodley and others, 2002).

Comments: Because of its large size, it has been popular in the aquaculture industry and its use has been encouraged by the U.S. Department of Agriculture. Culture in the United States began in Hawaii in the early 1970s (FAO 2005–2012). Generally, it is cultured in tropical and subtropical climates such as Texas, South Carolina, and Mississippi; however, it has been experimentally cultured in a temperate climate in Kentucky (Tidwell and others, 2005).

Procambarus hayi (Faxon, 1884) (Straightedge Crayfish)

Habitat: The Straightedge Crayfish is found in freshwater, semi-permanent to permanent streams; tolerant to sluggish flow (Payne, 1972).

Life History: The breeding season occurs from March through September when females lay eggs in burrows from August to November. Egg hatching occurs from 18 to 31 days after they are laid. Up to and including the fourth instar may remain attached to the female; third and fourth instars are the first juveniles to enter open water. Juveniles are considered adults after 6-8 months, reaching carapace length range of 36.0-45.5 U.S. Forest Service mm; adult carapace length reached 55.0 mm after 1 year. The adults remain in deeper water during the day, moving into shallow waters at night (Payne, 1972).

Native Range: Native to western drainages of the Tombigbee River from the Noxubee River in Mississippi and northward in the state into drainages of the Yalobusha, Skuna, Yocona, Tallahatchie, and Coldwater rivers (Payne, 1972). The home range is not depicted on the map below.

Nonindigenous Range: In 2009, this crayfish was documented for the first time in Georgia. It was collected from a roadside ditch in Newton County (Oconee drainage); it was later found in two Newton County streams also in Georgia (Skelton, 2010).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown

Vector: Unknown, but most likely a bait bucket release.

Impacts: Unknown, but could have the potential to threaten native species of crayfish.

Crustacean References Cited

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Avenant-Oldewage, A., and Swanepoel, J.H., 1993, The male reproductive system and mechanisms of sperm transfer in Argulus japonicus (Crustacea: Branchiura): Journal of Morphology, 215(1):51—63.

Barko, V.A., and Hrabik, R.A., 2004, Abundance of Ohio shrimp (Macrobrachium ohione) and glass shrimp (Palaemonetes kadiakensis) in the unimpounded upper Mississippi River: American Midland Naturalist, v. 151, no. 2, p. 265—273.

Bauer, R.T., and Delahoussaye, J., 2008, Life history migrations of the amphidromous river shrimp Macrobrachium ohione from a continental large river system: Journal of Crustacean Biology, v. 8, no. 4, p. 622—632.

Bonou, C.A., Pagano, M., and Saint-Jean, L., 1991, Développement et croissance en poids de Moina (cf) micrura et de Mesocyclops ogunnus dans un milieu saumâtre tropical—Les étangs de pisciculture de Layo (Côte-d’Ivoire): Revue d’Hydrobiologie Tropicale, v. 24, p. 287–303.

Bowles E.E., Aziz, K., and Knight, C.L., 2000, Macrobrachium (Decapoda: Caridea: Palaemonidae) in the contiguous United States—A review of the species and an assessment of threats to their survival: Journal of Crustacean Biology, v. 29, p. 158—171.

Bruno, M.C., Reid, J.W., and Perry, S.A., 2000, New records of copepods from Everglades National Park (Florida)—Description of two new species of Elaphoidella (Harpacticoida, Canthocamptidae), and supplementary description of Diacyclops nearcticus Kiefer (Cyclopoida, Cyclopidae): Crustaceana v. 73, p. 1,171—1,204.

Bunkley-Williams, L.; and Williams, E.H., Jr., 1994, Parasites of Puerto Rican freshwater sport fishes: San Juan, Puerto Rico, Puerto Rico Department of Natural and Environmental Resources, and Mayaguez, Puerto Rico, Department of Marine Sciences, University of Puerto Rico, 168 p.

Byron, E.R.; and Saunders, J.F., III., 1981, Colonization of Lake Tahoe, California, Nevada, USA and other western habitats by the copepod Skistodiaptomus pallidus (Calanoida): Southwestern Naturalist, v. 26, no. 1, p. 82—83.

Chapman, M.A., Green, J.D., and Northcote, T.G., 1985, Seasonal dynamics of Skistodiaptomus pallidus Herrick and other zooplankton populations in Deer Lake, S. W. British Columbia: Journal of Plankton Research, v. 7, no. 6, p. 867—876.

Duggan, I.C., Green, J.D., and Burger, D.F., 2006, First New Zealand records of three non-indigenous zooplankton species—Skistodiaptomus pallidus, Sinodiaptomus valkanovi, and Daphnia dentifera: New Zealand Journal of Marine and Freshwater Research, v. 40, p. 561—569.

Food and Agriculture Organization of the United Nations, 2005—2012, Cultured Aquatic Species Information Programme — Macrobrachium rosenbergii — Cultured Aquatic Species Information Programme, Text by New, M.B., IN: FAO Fisheries and Aquaculture Department [online], Rome, Updated 1 January 2004. Also available at http://www.fao.org/fishery/culturedspecies/Macrobrachium_rosenbergii/en Accessed 5/24/2012.

Geddes, M.C., and Cole, G.A., 1981, Variation in sexual size differentiation in North American diaptomids (Copepoda: Calanoida)—Does variation in the degree of dimorphism have ecological significance?: Limnology and Oceanography, v. 26, no. 2, p. 367—374.

Geiling, W.T., and Campbell, R.S., 1972, The effect of temperature on the development rate of the major life stages of Diaptomus pallidus Herrick: Limnology and Oceanography, v. 17, p. 304—307.

Gophen, M., 1977, Food and feeding habits of Mesocyclops leuckarti (Claus) in Lake Kinneret (Israel): Freshwater Biology, v. 7, p. 513–518.

Gresty, K.A., Boxshall, G.A., and Nagasawa, K., 1993, The fine structure and function of the cephalic appendages of the branchiuran parasite, Argulus japonicus Thiele: Philosophical Transactions of the Royal Society of London: Biological Sciences, v. 339, no. 1287, p. 119—135.

Hameed, A.S.S., Charles, M.X., and Anilkumar, M., 2000, Tolerance of Macrobrachium rosenbergii to white spot syndrome virus: Aquaculture, 183: 207—213.

Hobbs, H.H., and Lodge, D.M., 2010, Decapoda, IN: Thorp, J. H., and Covich, A.P., (eds.), Ecology and classification of North American freshwater invertebrates (3rd ed.): Amsterdam, Academic Press, p. 901—967.

Hribar, L.J., and Reid, J.W., 2008, New records of copepods (Crustacea) from the Florida Keys: Southeastern Naturalist, v. 7, no. 2, p. 219—228.

Hubschman, J.H., 1983, Diaptomus pallidus Herrick, 1879 (Crustacea: Copepoda) as an intermediate host for Tanaorhamphus longirostris (Van Cleave, 1913) (Acanthocephala: Neoechinorhynchidae): Journal of Parasitology, v. 69, no. 5, p. 930—932.

Hudson, P.L., and Lesko, L.T., 2003, Free-living and parasitic copepods of the Laurentian Great Lakes: Keys and details on individual species: Ann Arbor, Michigan, U.S. Geological Survey Great Lakes Science Center home page. (Also available at http://www.glsc.usgs.gov/greatlakescopepods/.)

Huner, J.V., 1977, Observations on the biology of the river shrimp from a commercial bait fishery near Port Allen, Louisiana, IN: Proceedings of the 31st Annual Conference of the Southeastern Association of Fish and Wildlife Agencies, 1977:380—386.

Karaytug, S., and Boxshall, G.A., 1998, Partial revision of Paracyclops Claus, 1893 (Copepoda, Cyclopoida, Cyclopidae) with descriptions of four new species: Bulletin of the Natural History Museum of London (Zoology), v. 64, p. 111—205.

Krieger, K.A., and Klarer, D.M., 1991, Zooplankton dynamics in a Great Lakes coastal marsh: Journal Great Lakes Research, v. 17, p. 255—269.

Kruger, I., van As, J.G., and Saayman, J.E., 1983, Observations on the occurrence of the fish louse Argulus japonicus Thiele, 1900 in the western Transvaal: South African Journal of Zoology, v. 18, no. 4, p. 408— 410.

Marten, G.G., and Reid, J.W., 2007, Cyclopoid copepods, IN: Floore, T.G. (ed.), Biorational control of mosquitoes. American Mosquito Control Association Bulletin 7: Supplement to the Journal of the American Mosquito Control Association, v. 23, no. 2, p. 65–92.

Mather, P.B., and de Bruyn, M., 2003, Genetic diversity in wild stocks of the giant freshwater prawn (Macrobrachium rosenbergii): implications for aquaculture and conservation: Naja, v. 26, no. 4, p. 4—7.

Matsumura-Tundisi, T., and Silva, W. M., 2002, Occurence of Mesocyclops ogunnus Onabamiro, 1957 (Copepoda Cyclopoida) in water bodies of Sao Paulo State, identified as Mesocyclops kieferi Van de Velde, 1984: Brazilian Journal of Biology, v. 62, no. 4A, p. 615—620.

McLaughlin, P.L., and 39 others, 2005, Common and scientific names of aquatic invertebrates from the United States and Canada—Crustaceans: American Fisheries Society Special Publication 31, Bethesda, Maryland, American Fisheries Society, 545 p.

Micieli, M.V., Garcia, J.J., and Becnel, J.J., 2000, Life cycle and description of Amblyospora camposi n. sp. (Microsproidia: Amblyosporidae) in the mosquito Culex renatoi (Diptera, Cluicidae) and copepod Paracyclops fimbriatus fimbriatus (Copepoda, Cyclopidae): Journal of Eukaryot Microbiology, v. 47, no. 6, p. 575-580.

Mills, E.L., Leach, J.H., Carlton, J.T., and Secor, C.L., 1993, Exotic species in the Great Lakes—A history of biotic crises and anthropogenic introductions: Journal of Great Lakes Research, v. 19, no 1, p. 1—54.

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Shafir, A., and Van As, J.G., 1986, Laying, development of hatching of eggs of the fish ectoparasite Argulus japonicus (Crustacea: Branchiura): Journal of Zoology (London), v. 210, p. 401—414.

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Torke, B., 2001, The distribution of calanoid copepods in the plankton of Wisconsin Lakes: Hydrobiologia, v. 453—454, p. 351—365.

Williams, E.H., Jr.; Bunkley-Williams, L.; Lilyestrom, C.G., and Ortiz-Corps, E.A.R., 2001, A review of recent introductions of aquatic invertebrates in Puerto Rico and implications for the management of nonindigenous species: Caribbean Journal of Science, v. 37, no. 3—4, p. 246—251.

Williamson, C.E., and Butler, N.M., 1986, Predation on rotifers by the suspension-feeding calanoid copepod Diaptomus pallidus: Limnology and Oceanography, v. 31, p. 393—402.

Williamson, C.E., and Vanderploeg, H.A., 1988, Predatory suspension-feeding in Diaptomus—Prey defenses and the avoidance of cannibalism: Bulletin of Marine Science, v. 43, no. 3, p. 561—572.

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Woodley, C.M., Slack, W.T., Peterson, M.S., and Vervaeke, W.C., 2002, Occurrence of the nonindigenous giant Malaysian prawn, Macrobrachium rosenbergii (De Man, 1879) in Simmons Bayou, Mississippi, U.S.A.: Crustaceana, v. 75, no. 8, p. 1025—1031.

Wynne, F., 2000, Grow-out culture of freshwater prawns in Kentucky: Mayfield, Kentucky, Kentucky State University Cooperative Extension Program. Also available at: http://web.archive.org/web/20080821114126/http://www.aquanic.org/publicat/state/ky/prawn_ext.htm Accessed 5/25/2012.

MOLLUSKS

Overview

There have been 22 species of aquatic mollusks introduced to Region 4 as early as the 1940s (Figure 8; Table 3; Table 4). Gastropods, or snails, make up the majority (18) of these introductions followed by bivalves (4), also referred to as clams or mussels (Figure 8). The origins of these species are primarily from other continents (Figure 9). Since 2000, there have been only four newly documented mollusk species in Region 4 (Table 3).

Snails Bivalves

Figure 8. Total number of introduced mollusk species by taxonomic group.

3 Asia/Eurasia

10 South and Central America North America 9

Figure 9. Total number of introduced mollusk species by continental origin.

Among mollusks, snails undoubtedly have the greatest number of species introductions because of their popularity in the aquarium trade, water garden, and ornamental plant industries. All 15 species designated as “Aquarium releases” are snails (Figure 10). Aside from what we find as more typical introductions, oddly misplaced organisms continued to be found in open waters. For example, in 2003 in Arkansas’s Lake Conway, a live octopus (Octopus vulgaris) was pulled out of the water as it was clinging to a gate at the dam. An incident such as this is mostly the result of an aquarium release and is illegal in this particular state. Because this particular introduction would never result in an established population, the octopus was not treated in this report with a species account.

1 Aquarium release 3

Unknown

3 Shipping

15 Escaped aquaculture

Figure 10. Total number of introduced mollusk species by vector.

Number of New Species

10 The number of new species 8 documented since 2000 has the 6 potential to surpass the number introduced in previous 25-year 4 period (Figure 11). 2

0 1900-1925 1926-1950 1951-1975 1976-2000 2001-2012

Figure 11. Total number of mollusk species introduced in 25-year intervals. (Note that the current interval is only 12 years.)

Species of concern in the region are Quagga Mussels because of the ecological harm visible in the Great Lakes and mostly likely soon to be seen in western states, and applesnails which can devastate agricultural crops

Table 3. All nonindigenous mollusk species introductions into U.S. Fish and Wildlife Service Region 4. Group Family Scientific Name Common Name First Continent of Origin Likely Vector Population Year Status Bivalves Corbiculidae Corbicula fluminea Asian Clam 1957 Asia unknown Established Bivalves Dreissenidae Dreissena bugensis Quagga Mussel 2004 Eurasia shipping Unknown Bivalves Dreissenidae Dreissena polymorpha † Zebra Mussel 1991 Eurasia shipping Established Bivalves Sphaeriidae Pisidium punctiferum † Striate Peaclam 1999 South/Central America unknown Established Gastropods Ampullariidae Marisa cornuarietis Giant Rams-horn Snail 1957 South America released aquarium Established Gastropods Ampullariidae Pomacea bridgesi Spiketop Applesnail 1966 South America released aquarium Established Gastropods Ampullariidae Pomacea canaliculata † Channeled Applesnail 1992 South America released aquarium Established Gastropods Ampullariidae Pomacea cumingii applesnail 1954 South America released aquarium Established Gastropods Ampullariidae Pomacea diffusa † Spike-topped Applesnail 1966 South America released aquarium Unknown Gastropods Ampullariidae Pomacea haustrum † Titan Applesnail 1978 South America released aquarium Established Gastropods Ampullariidae Pomacea maculata Giant Applesnail 1989 South America escaped captivity aquaculture Established Gastropods Ampullariidae Pomacea paludosa Florida Applesnail 1970 North America unknown Established Gastropods Lymnaeidae Radix auricularia European Ear Snail 1969 Europe released aquarium Unknown Gastropods Planorbidae Biomphalaria glabrata Bloodfluke Planorb 1995 South America shipping Established Gastropods Thiaridae Melanoides tuberculata Red-rim Melania 1970 Asia released aquarium Established Gastropods Thiaridae Melanoides turriculus Fawn Melania 1995 Asia released aquarium Established Gastropods Thiaridae Tarebia granifera Quilted Melania 1949 Asia released aquarium Established Gastropods Thiaridae Thiara scabra * Pagoda Tiara 2006 Asia released aquarium Unknown Gastropods Viviparidae Cipangopaludina chinensis malleata Chinese Mysterysnail 1965 Asia released aquarium Established Gastropods Viviparidae Cipangopaludina japonica † Japanese Mysterysnail 1994 Asia released aquarium Unknown Gastropods Viviparidae Viviparus georgianus Banded Mysterysnail 2008 North America released aquarium Established Gastropods Viviparidae Viviparus subpurpureus Olive Mysterysnail 2008 North America released aquarium Established Bold = New to U.S. Fish and Wildlife Service Region 4 since 2000 * = New to Country † = Introduced to USFWS Region 4 prior to 2001 but not included in last report

Table 4. All nonindigenous mollusk species introductions into U.S. Fish and Wildlife Service Region 4 by state/territory. Group Family Scientific Name Common Name AL AR FL GA KY LA MS NC PR SC TN VI Bivalves Corbiculidae Corbicula fluminea Asian Clam X X X X X X X X X X X Bivalves Dreissenidae Dreissena bugensis Quagga Mussel N Bivalves Dreissenidae Dreissena polymorpha † Zebra Mussel X X X X X X Bivalves Sphaeriidae Pisidium punctiferum † Striate Peaclam X Gastropods Ampullariidae Marisa cornuarietis Giant Rams-horn Snail X N Gastropods Ampullariidae Pomacea bridgesi Spiketop Applesnail X Gastropods Ampullariidae Pomacea canaliculata † Channeled Applesnail N X Gastropods Ampullariidae Pomacea cumingii applesnail X Gastropods Ampullariidae Pomacea diffusa † Spike-topped Applesnail N X Gastropods Ampullariidae Pomacea haustrum † Titan Applesnail X Gastropods Ampullariidae Pomacea maculata Giant Applesnail N X N N N N N Gastropods Ampullariidae Pomacea paludosa Florida Applesnail X X X X Gastropods Lymnaeidae Radix auricularia European Ear Snail X Gastropods Planorbidae Biomphalaria glabrata Bloodfluke Planorb X Gastropods Thiaridae Melanoides tuberculata Red-rim Melania X X N N X Gastropods Thiaridae Melanoides turriculus Fawn Melania X Gastropods Thiaridae Tarebia granifera Quilted Melania X N X Gastropods Thiaridae Thiara scabra * Pagoda Tiara N Gastropods Viviparidae Cipangopaludina chinensis malleata Chinese Mysterysnail X X Gastropods Viviparidae Cipangopaludina japonica † Japanese Mysterysnail N X N X Gastropods Viviparidae Viviparus georgianus Banded Mysterysnail N Gastropods Viviparidae Viviparus subpurpureus Olive Mysterysnail N Bold = New to U.S. Fish and Wildlife Service Region 4 since 2000 * = New to United States † = Introduced to USFWS Region 4 prior to 2001 but not included in last report X = Present in or prior to 2000 N = New since 2000

Species Accounts of Mollusks New to Region 4

Dreissena bugensis Andrusov, 1897 (Quagga Mussel)

Habitat: Quagga Mussels are found in freshwater rivers, lakes, and reservoirs. In North American populations, they are not known to tolerate salinities greater than 5 ppt (Spidle and others, 1995). Optimal temperature for normal function is generally below 25 oC, but as warm water populations acclimate, the upper thermal limit could increase. Water temperatures of 28 oC begin to cause significant mortality, and U.S. Geological Survey temperatures of 32–35 oC are considered lethal for Dreissena species (Antonov and Shkorbatov, 1990 as cited in Mills, and others, 1996). The depth at which the mussels live varies depending on water temperature. They generally are not found in lakes near the shore in shallow water due to wave action. The Quagga Mussel can inhabit hard and soft substrates, including sand and mud, down to depths of at least 130 meters (m) (Mills and others, 1993).

Life History: A small bivalve mussel averaging 30 mm in length, the shell is somewhat D-shaped and most often has a striped pattern (Morton, 1969). Females lay thousands of eggs after the water warms in the spring to generally above 12 oC with a few exceptions (Nichols, 1996). These barely visible eggs are then fertilized by the males in the open water where they hatch and become free-swimming larvae (Ram and others, 1996). The larvae grow a velum used for feeding and swimming, and then is known as a veliger. After a month or more from hatching, they settle onto hard substrates and attach themselves using an organ called a byssus (Mackie and others, 1989). From this postveliger stage they grow to reach the juvenile and adult stages and are sexually mature by their second year (Morton, 1969). As a filter- feeder, they siphon algae, detritus, bacteria, and other small organisms from the water for food. The lifespan of a Quagga Mussel is about 2–5 years (Mackie and others, 1990).

Native Range: Dreissena bugensis is indigenous to the Dneiper River drainage of Ukraine and Ponto- Caspian Sea. It was discovered in the Bug River in the Ukraine in 1890 by Andrusov, who named the species in 1897 (Mills and others, 1996).

Nonindigenous Range: In 2004, very limited numbers of Quagga Mussels were found in just two of many sample sites on the Ohio River in Kentucky (Grigorovich and others, 2008). Other locations where Quagga Mussels have become well established are in the Great Lakes and more recently in 2007 in the lower Colorado River, including Lake Mead in Nevada.

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: The introduction of D. bugensis into the Great Lakes appears to be the result of ballast water discharge from transoceanic ships that were carrying veligers, juveniles, or adult mussels. The genus Dreissena is highly polymorphic and prolific with high potential for rapid adaptation attributing to its rapid expansion and colonization (Mills and others, 1996). Still, there are other factors that can aid in the spread of this species across North American waters, such as larval drift in river systems or fishing and boating activities that allow for overland transport or movement between water basins.

Impacts: Quagga mussels are prodigious water filterers, removing substantial amounts of phytoplankton and suspended particulate from the water. As such, their impacts are similar to those of the Zebra Mussel (Dreissena polymorpha). By removing the phytoplankton, Quagga Mussels decrease the food source for zooplankton, therefore altering the food web. Impacts associated with the filtration of water include increases in water transparency, decreases in mean chlorophyll a concentrations, and accumulation of pseudofeces (Claxton and others, 1998). Water clarity increases light penetration, causing a proliferation of aquatic plants that can change species dominance and alter the entire ecosystem. The pseudofeces that are produced from filtering the water accumulate, creating a foul environment. As the waste particles decompose, oxygen is used up, and the pH becomes very acidic and toxic byproducts are produced. In addition, Quagga Mussels accumulate organic pollutants within their tissues to levels more than 300,000 times greater than concentrations in the environment and these pollutants are found in their pseudofeces, which can be passed up the food chain, therefore increasing wildlife exposure to organic pollutants (Snyder and others, 1997). Dreissena species ability to rapidly colonize hard surfaces causes serious economic problems. These major biofouling organisms can clog water intake structures, such as pipes and screens, therefore reducing pumping capabilities for power and water treatment plants, costing industries, companies, and communities. Recreation-based industries and activities also have been impacted where docks, breakwalls, buoys, boats, and beaches have all been heavily colonized. Many of the potential impacts of Dreissena are unclear due to the limited time scale of North American colonization.

Thiara scabra (Müller, 1774) (Pagoda Tiara)

Habitat: This snail lives in freshwater streams (Maciolek and Ford, 1987) and coastal brackish waters (Sri-aroon and others, 2004).

Life History: This species is a small snail at 20 mm in shell height with 6–8 whorls (Thompson and others, 2009). Reproduction is parthenogenic (Thompson and others, 2009). Embryos develop in a brood pouch to a shelled juvenile and are fed by a yolk sac and a placenta-like epithelium in the brood pouch (Glaubrecht, 2006). D. Strom They have few young, but are well developed when they hatch (Glaubrecht, 2006).

Native Range: South and Southeast Asia and Indo-Australian Archipelago and western Pacific Islands (Brandt, 1974 as cited in Thompson and others, 2009).

Nonindigenous Range: Thiara scabra has been found on the east edge of Lake Okeechobee in two Florida counties in 2006 and 2007 (Thompson and others, 2009).

Impacts: Unknown; however, this species is susceptible, albeit at a low rate of 1.2 percent, to trematode infections in Thailand (Dechruksa and others, 2007).

Comments: Because of its mode of reproduction, a single specimen can start a population (Thompson and others, 2009). It is not known if these populations persist in Florida. It was reported that none were found at one previously infested site in 2008.

Viviparus georgianus (Lea, 1934) (Banded Mysterysnail)

Habitat: The Banded Mysterysnail is generally found in freshwater slow- moving streams, ponds, lakes, and marshes (Duch, 1976).

Life History: The sexes are separate, and females give birth to live young. A large snail 20–45 mm in height, with up to 4 dark bands (Thompson, 2004), 4– 5 whorls, and a broadly globose conic shape (Burch and Tottenham, 1980). They graze on algae and diatoms (Duch, 1976) as well as fish eggs and fish embryos (Eckblad and Shealy, 1972).

Native Range: Widespread in Florida, Georgia, northern Alabama, Louisiana, U.S. Geological Survey and north mainly in the Mississippi River system to Illinois, northwestern Indiana (Clench, 1962). The home range is not depicted on the map below.

Nonindigenous Range: The Banded Mysterysnail has been found in the Lake Marion and Lake Moultrie area of South Carolina (Dillon and others, 2006), and in many lakes in the Great Lakes region and New England (Clench, 1962).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: It is difficult to determine the pathway of introduction for native species not known to be in the aquarium trade; however, because the native range is relatively close, these reservoir introductions most likely are the result of unintentional release of water containing small snails through recreational activities.

Impacts: Unknown.

Comments: Collections of this snail outside its native range date back to the early 1900s (Clench, 1962).

Viviparus subpurpureus (Say, 1829) (Olive Mysterysnail)

Habitat: The Olive Mysterysnail inhabits freshwater streams, ponds, lakes, and marshes.

Life History: The sexes are separate and the females give birth to live young. A large snail with a broadly globose conic shape, which may have 3 dark colored bands but not always (Burch and Tottenham, 1980), and a 33-mm shell height (Clench and Fuller, 1965).

Native Range: Mississippi River drainage from Iowa and Illinois to Louisiana, and David Knott in the Gulf drainage in east Texas, west Louisiana, and east Mississippi (Clench and Fuller, 1965). The home range is not depicted on the map below.

Nonindigenous Range: The Olive Mysterysnail has been found in the Lake Marion and Lake Moultrie area of South Carolina (Dillon and others, 2006).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: It is difficult to determine the pathway of introduction for native species not known to be in the aquarium trade. Because the native range is relatively close, these reservoir introductions most likely are the result of unintentional release of water containing small snails through recreational activities.

Impacts: Unknown.

Mollusk References Cited

Antonov, P.I., and Shkorbatov, G.L., 1990, Ecological-physiological characteristics of Dreissena of the Lower Reaches of the Dnieper River. In Vid i yego areale. Biologiya, ekologiya i produktivnosf vodnykh bespozvonochnykh (Species in its distribution range. Biology, ecology and production of aquatic invertebrates); p. 126—130. Minsk. (In Russian).

Brandt, R.A.M., 1974, The non-marine aquatic Mollusca of Thailand: Archiv für Molluskenkunde, v. 105, p. 1—423.

Burch, J.B., and Tottenham, J.L., 1980, North American freshwater snails—Species lists, ranges and illustrations: Walkerana, v. 1, n. 3, p. 81—215.

Claxton, W.T., Wilson, A.B., Mackie, G.L., and Boulding, E.G., 1998, A genetic and morphological comparison of shallow- and deep-water populations of the introduced dreissenid bivalve Dreissena bugensis: Canadian Journal of Zoology, v. 76, n. 7, p. 1,269—1,276.

Clench, W.J., 1962, A catalogue of the Viviparidae of North America with notes on the distribution of Viviparus georgianus: Occasional Papers on Mollusks, v. 2, p. 261—287.

Clench, W.J., and Fuller, S.L.H., 1965, The genus Viviparus (Viviparidae) in North America: Occasional Papers on Mollusks, v. 2, no. 32, p. 385—412.

Dechruksa, W., Krailas, D., Ukong, S., Inkapatanakul, W., and Koonchornboon, T., 2007, Trematode infections of the freshwater snail family Thiaridae in the Khek River, Thailand: Southeast Asian Journal of Tropical Medicine and Public Health, v. 38, no. 6, p. 1,016—1,028.

Dillon, R.T., Jr.; Watson, B.T.; Stewart, T.W.; and Reeves, W.K., 2006, The freshwater gastropods of North America. Also available at: http://www.fwgna.org/FWGSC/viviparus.pdf

Duch, T.M., 1976, Aspects of the feeding habits of Viviparus georgianus: The Nautilus, v. 90, no. 1, p. 7— 10.

Eckblad, J.W.; and Shealy, M.H., Jr., 1972, Predation on largemouth bass embryos by the pond snail Viviparus georgianus: Transactions of the American Fisheries Society, v. 101, no. 4, p. 734—738.

Glaubrecht, M., 2006, Independent evolution of reproductive modes in viviparous freshwater Cerithioidea (Gastropoda, Sorbeoconcha)—A brief review: Basteria Supplement, v. 3, p. 23—28.

Grigorovich, I.A., Angradi, T.R., and Stepien, C.A., 2008, Occurrence of the quagga mussel (Dreissena bugensis) and the zebra mussel (Dreissena polymorpha) in the Upper Mississippi River System: Journal of Freshwater Ecology, v. 23, no. 3, p. 429—435.

Maciolek, J.A., and Ford, J.I., 1987, Macrofauna and environment of the Nanpil-Kiepw River, Ponape, Eastern Caroline Islands: Bulletin of Marine Science, v. 41, no. 2, p. 623—632.

Mackie, G.L., Gibbons, W.N., Muncaster, B.W., and Gray, I.M., 1989, The zebra mussel, Dreissena polymorpha—A synthesis of European experiences and a preview for North America, Report prepared for Ontario Ministry of Natural Resources. Queen's Printer for Ontario, [ISBN 0-7729-5647-2].

Mills, E.L., Dermott, R.M., Roseman, E.F., Dustin, D., Mellina, E., Conn, D.B., and Spidle, A.P., 1993, Colonization, ecology, and population structure of the quagga mussel (Bivalvia: Dreissenidae) in the lower Great Lakes: Canadian Journal of Fisheries and Aquatic Sciences, v. 50, p. 2,305—2,314.

Mills, E.L., Rosenberg, G., Spidle, A.P., Ludyanskiy, M., Pligin, Y., and May, B., 1996, A review of the biology and ecology of the quagga mussel (Dreissena bugensis), a second species of freshwater dreissenid introduced to North America: American Zoology, v. 36, p. 271—286.

Morton, B., 1969, Studies on the biology of Dreissena polymorpha Pall. I. General anatomy and morphology: Proceedings of the Malacological Society of London, v. 38, p. 301—321.

Nichols, S.J., 1996, Variations in the reproductive cycle of Dreissena polymorpha in Europe, Russia, and North America: American Zoologist, v. 36, no. 3, p. 311—325.

Ram, J.L., Fong, P.P., and Garton, D.W., 1996, Physiological aspects of zebra mussel reproduction: maturation, spawning, and fertilization: American Zoologist, v. 36, no. 3, p. 326—338.

Snyder, F.L., Hillgendorf, M.B., and Garton, D.W., 1997, Zebra mussels in North America, the invasion and its implications: Ohio State Sea Grant OHSU-FS-045, Columbus, Ohio. 4 p.

Spidle, A.P., Mills, E.L., and May, B., 1995, Limits to tolerance of temperature and salinity in the quagga mussel (Dreissena bugensis) and the zebra mussel (Dreissena polymorpha): Canadian Journal of Fisheries and Aquatic Sciences, v. 52, p. 2,108—2,119.

Sri-aroon, P., Lohachit, C., and Harada, M., 2004, Survey of brackish-water snails in eastern Thailand: Southeast Asian Journal of Tropical Medicine and Public Health, v. 35(Suppl. 1), p. 151—155.

Thompson, F.G., 2004, An identification manual for the freshwater snails of Florida: Gainesville, Florida, Florida Museum of Natural History, University of Florida. Also available at: http://www.flmnh.ufl.edu/malacology/fl-snail/snails1.htm Accessed 1/7/2012.

Thompson, F.G., Heyn, M.W., and Campbell, D.N., 2009, Thiara scabra (O. F. Mueller, 1774)—The introduction of another Asian freshwater snail into the United States: The Nautilus v. 123, no. 1, p. 21— 22.

AMPHIBIANS AND REPTILES

Overview

A total of 18 new species have been added to the region since 2000 (Table 5; Table 6): 7 amphibians and 11 reptiles. Nearly all reports are from Krysko and others (2011). Amphibians and reptiles were not included in the 2001 Region 4 Report, so this section is unique to this update.

Turtles and frogs are the most numerous groups of introduced aquatic herptiles, followed by salamanders, crocodilians, and snakes (Figure 12).

A B 2

Crocodilians 3 Frogs Snakes Salamanders 6 4 3 Turtles

Figure 12. Total number of introduced amphibian (A) and reptile (B) species by taxonomic group.

Most newly introduced aquatic herptiles originate from the Western hemisphere (South and Central American and North American transplants), followed by Asia and Eurasia (Figure 13).

South and A B South and 1 1 Central America 1 Central America North America 1 5 North America Asia/Eurasia 5 Asia/Eurasia Africa 4

Figure 13. Number of new (since 2000) introduced amphibian (A) and reptile (B) species by continental origin.

Pet release accounts for 100 percent of all new amphibian and reptile species introductions (Figure 14).

Twelve amphibian and reptile species never introduced in the country before have been found since 2000. Of these, 11 were from foreign countries and 1 was a native transplant (Table 5).

A B

Pet release Pet release

7 11

Figure 14. Number of new (since 2000) introduced amphibian (A) and reptile (B) species by vector.

Only the Yellow Anaconda has been found in more than one state since 2001. In this case, they are separate introductions rather than a species that is dispersing.

Table 5. All nonindigenous amphibian and reptile introductions into U.S. Fish and Wildlife Service Region 4. First Population Group Family Name Common Name Year Continent of Origin Likely Vector Status Frogs Bombinatoridae Bombina orientalis * Oriental Fire-bellied Toad 2007 Eurasia released pet unknown Frogs Bufonidae Atelopus zeteki Panamanian Golden Frog 1963 South America released pet failed Frogs Bufonidae Incilius nebulifer Coastal Plain Toad 1883 North America unknown established Frogs Bufonidae Rhaebo blombergi Columbian Giant Toad 1963 South America released pet failed Frogs Bufonidae Rhinella marina Cane Toad 1935 North and South America stocked for biocontrol established Frogs Eleutherodactylidae Eleutherodactylus coqui Coqui 1973 Caribbean hitchhiker on plants established Frogs Eleutherodactylidae Eleutherodactylus cystignathoides Rio Grande Chirping Frog 1998 North America hitchhiker on plants established Frogs Eleutherodactylidae Eleutherodactylus planirostris Greenhouse Frog 1863 Caribbean hitchhiker on plants established Frogs Eleutherodactylidae Eleutherodactylus portoricensis Forest Coqui 1964 Caribbean hitchhiker on plants failed Frogs Hylidae Hyla cinerea Green Treefrog 1975 North America hitchhiker on plants established Frogs Hylidae Litoria caerulea Australian Green Treefrog 1999 Australia released pet established Frogs Hylidae Osteopilus septentrionalis Cuban Treefrog 1931 Caribbean hitchhiker on plants established Frogs Hylidae Pachymedusa dacnicolor Mexican Leaf Frog 1964 South America released pet failed Frogs Hylidae Pseudacris sierra Sierran Chorus Frog 1983 North America hitchhiker unknown Frogs Hylidae Scinax rubra Common Scinax 1998 North America hitchhiker established Frogs Hyperoliidae Afrixalus fornasini * Fornasini's Spiny Reed Frog 2010 Africa released pet unknown Frogs Microhylidae Kaloula pulchra * Malaysian Painted Frog 2006 Asia released pet unknown Frogs Pipidae Hymenochirus boettgeri Zaire Dwarf Clawed Frog 1964 Africa released pet failed Frogs Pipidae Pipa pipa * Suriname Toad 2007 South America released pet unknown Frogs Pipidae Xenopus laevis African Clawed Frog 1964 Africa released pet failed Frogs Ranidae Lithobates catesbeianus American Bullfrog 1967 North America stocked for food established Frogs Ranidae Lithobates grylio Pig Frog 1998 North America stocked for food established Salamanders Amphiumidae Amphiuma tridactylum * Three-toed Amphiuma 2009 North America released pet unknown Salamanders Plethodontidae Desmognathus quadramaculatus Black-bellied Salamander 1991 North America released bait established Salamanders Plethodontidae Desmognathus santeetlah Santeetlah Dusky Salamander 1996 North America released bait unknown Salamanders Salamandridae Cynops orientalis * Oriental Fire-bellied Newt 2009 Asia released pet unknown Japanese Fire-bellied Salamanders Salamandridae Cynops pyrrhogaster Salamander 1964 Asia released pet failed

Table 5. All nonindigenous amphibian and reptile introductions into U.S. Fish and Wildlife Service Region 4. (Continued) First Population Group Family Name Common Name Year Continent of Origin Likely Vector Status Notophthalmus viridescens Salamanders Salamandridae viridescens Red-spotted Newt 1964 North America released pet failed Salamanders Salamandridae Pachytriton labiatus * Paddle-Tail Newt 2010 Asia released pet unknown Crocodilians Alligatoridae Alligator mississippiensis American Alligator 1995 North America released pet failed Crocodilians Alligatoridae Caiman crocodilus Common Caiman 1959 South America released pet established Cuvier's Smooth-fronted Crocodilians Alligatoridae Paleosuchus palpebrosus * Caiman 2008 South America released pet unknown Schneider's Smooth-fronted Crocodilians Alligatoridae Paleosuchus trigonatus Caiman 1999 South America released pet failed Crocodilians Crocodylidae Crocodylus acutus American Crocodile 2004 North America escaped captivity failed African Slender-snouted Crocodilians Crocodylidae Crocodylus cataphractus Crocodile 1986 Africa released pet failed Crocodilians Crocodylidae Crocodylus niloticus Nile Crocodile 1998 Africa escaped captivity aquaculture failed Snakes Acrochoridae Acrochordus javanicus Javan Filesnake 1970 Asia released pet failed Snakes Boidae Eunectes murinus * Green Anaconda 2004 South America released pet unknown Snakes Boidae Eunectes notaeus * Yellow Anaconda 2004 South America released pet unknown Snakes Colubridae Erpeton tentaculatus * Tentacled Snake 2010 Asia released pet failed Snakes Colubridae Fordonia leucobalia Crab-eating Water Snake 1971 Asia unknown failed Turtles Bataguridae Rhinoclemmys pulcherrima Painted Wood Turtle 2006 South America released pet unknown Turtles Bataguridae Rhinoclemmys punctularia Spot-legged Turtle 1996 South America released pet established Turtles Chelidae Chelus fimbriatus Matamata 1963 South America released pet failed Turtles Chelidae Platemys platycephala Twist-necked Turtle 1985 South America released pet failed Turtles Chelydridae Chelydra serpentina Snapping Turtle 2006 North America released pet unknown Turtles Chelydridae Macrochelys temminckii Alligator Snapping Turtle 2007 North America released pet unknown Turtles Emydidae Chrysemys picta Painted Turtle 1939 North America released pet unknown Turtles Emydidae Chrysemys picta bellii Western Painted Turtle 1933 North America released pet unknown Turtles Emydidae Chrysemys picta dorsalis Southern Painted Turtle 1964 North America released pet failed Turtles Emydidae Deirochelys reticularia chrysea Florida Chicken Turtle 1989 North America released pet unknown Turtles Emydidae Glyptemys insculpta Wood Turtle 2009 North America released pet unknown Turtles Emydidae Graptemys barbouri Barbour's Map Turtle 1993 North America released pet unknown

Table 5. All nonindigenous amphibian and reptile introductions into U.S. Fish and Wildlife Service Region 4. (Continued) First Population Group Family Name Common Name Year Continent of Origin Likely Vector Status Turtles Emydidae Graptemys ouachitensis Ouachita Map Turtle 1992 North America released pet unknown Turtles Emydidae Graptemys pseudogeographica False Map Turtle 2000 North America released pet unknown Turtles Emydidae Graptemys pseudogeographica kohnii Mississippi Map Turtle 1964 North America released pet failed Turtles Emydidae Trachemys dorbigni Brazilian Slider 1964 South America released pet failed Turtles Emydidae Trachemys scripta Pond Slider 1963 North America released pet failed Turtles Emydidae Trachemys scripta callirostris Columbian slider 1964 South America released pet failed Turtles Emydidae Trachemys scripta elegans Red-eared Slider 1958 North America released pet established Turtles Emydidae Trachemys scripta scripta Yellow-bellied Slider 1976 North America unknown unknown Turtles Emydidae Trachemys stejnegeri malonei Inagua Slider 1946 Caribbean stocked for food failed Turtles Kinosternidae Kinosternon scorpioides Scorpion Mud Turtle 1964 South America released pet failed Pacific Coast Giant Musk Turtles Kinosternidae Staurotypus salvinii * Turtle 2010 Central America released pet unknown Turtles Kinosternidae Sternotherus carinatus Razor-backed Must Turtle 1937 North America released pet unknown Turtles Pelomedusidae Pelusios subniger East African Black Mud Turtle 1999 Africa released pet failed Turtles Pelomedusidae Podocnemis lewyana Magdalena River Turtle 1964 South America released pet failed Turtles Pelomedusidae Podocnemis sextuberculata Six-tubercled River Turtle 1964 South America released pet failed Turtles Pelomedusidae Podocnemis unifilis Yellow-spotted River Turtle 1964 South America released pet failed Turtles Trionychidae Apalone ferox Florida Softshell 1900 North America released pet unknown Turtles Trionychidae Apalone spinifera Spiny Softshell 2008 North America released pet unknown Bold = New to U.S. Fish and Wildlife Service Region 4 since 2000 * = New to United States

Table 6. All nonindigenous amphibian and reptile introductions into U.S. Fish and Wildlife Service Region 4 by state/territory. Group Family Name Common Name AL AR FL GA KY LA MS NC PR SC TN VI Frogs Bombinatoridae Bombina orientalis * Oriental Fire-bellied Toad N Frogs Bufonidae Atelopus zeteki Panamanian Golden Frog X Frogs Bufonidae Incilius nebulifer Coastal Plain Toad X Frogs Bufonidae Rhaebo blombergi Columbian Giant Toad X Frogs Bufonidae Rhinella marina Cane Toad X X X X Frogs Eleutherodactylidae Eleutherodactylus coqui Coqui X X Frogs Eleutherodactylidae Eleutherodactylus cystignathoides Rio Grande Chirping Frog X Frogs Eleutherodactylidae Eleutherodactylus planirostris Greenhouse Frog X X X X X Frogs Eleutherodactylidae Eleutherodactylus portoricensis Forest Coqui X Frogs Hylidae Hyla cinerea Green Treefrog X Frogs Hylidae Litoria caerulea Australian Green Treefrog X Frogs Hylidae Osteopilus septentrionalis Cuban Treefrog X N X X Frogs Hylidae Pachymedusa dacnicolor Mexican Leaf Frog X Frogs Hylidae Pseudacris sierra Sierran Chorus Frog X Frogs Hylidae Scinax rubra Common Scinax X Frogs Hyperoliidae Afrixalus fornasini * Fornasini's Spiny Reed Frog N Frogs Microhylidae Kaloula pulchra * Malaysian Painted Frog N Frogs Pipidae Hymenochirus boettgeri Zaire Dwarf Clawed Frog X Frogs Pipidae Pipa pipa * Suriname Toad N Frogs Pipidae Xenopus laevis African Clawed Frog X X Frogs Ranidae Lithobates catesbeianus American Bullfrog X Frogs Ranidae Lithobates grylio Pig Frog X Salamanders Amphiumidae Amphiuma tridactylum * Three-toed Amphiuma N Salamanders Plethodontidae Desmognathus quadramaculatus Black-bellied Salamander X Salamanders Plethodontidae Desmognathus santeetlah Santeetlah Dusky Salamander X Salamanders Salamandridae Cynops orientalis * Oriental Fire-bellied Newt N Salamanders Salamandridae Cynops pyrrhogaster Japanese Fire-bellied Salamander X Salamanders Salamandridae Notophthalmus viridescens viridescens Red-spotted Newt X

Table 6. All nonindigenous amphibian and reptile introductions into U.S. Fish and Wildlife Service Region 4 by state/territory. (Continued) Group Family Name Common Name AL AR FL GA KY LA MS NC PR SC TN VI Salamanders Salamandridae Pachytriton labiatus * Paddle-Tail Newt N Crocodilians Alligatoridae Alligator mississippiensis American Alligator N N X N Crocodilians Alligatoridae Caiman crocodilus Common Caiman X X Crocodilians Alligatoridae Paleosuchus palpebrosus * Cuvier's Smooth-fronted Caiman N Schneider's Smooth-fronted Crocodilians Alligatoridae Paleosuchus trigonatus Caiman X Crocodilians Crocodylidae Crocodylus acutus American Crocodile N African Slender-snouted Crocodilians Crocodylidae Crocodylus cataphractus Crocodile X Crocodilians Crocodylidae Crocodylus niloticus Nile Crocodile N X Snakes Acrochoridae Acrochordus javanicus Javan Filesnake X Snakes Boidae Eunectes murinus * Green Anaconda N Snakes Boidae Eunectes notaeus * Yellow Anaconda N N Snakes Colubridae Erpeton tentaculatus * Tentacled Snake N Snakes Colubridae Fordonia leucobalia Crab-eating Water Snake X Turtles Bataguridae Rhinoclemmys pulcherrima Painted Wood Turtle N Turtles Bataguridae Rhinoclemmys punctularia Spot-legged Turtle X Turtles Chelidae Chelus fimbriatus Matamata X Turtles Chelidae Platemys platycephala Twist-necked Turtle X Turtles Chelydridae Chelydra serpentina Snapping Turtle N Turtles Chelydridae Macrochelys temminckii Alligator Snapping Turtle N Turtles Emydidae Chrysemys picta Painted Turtle X X Turtles Emydidae Chrysemys picta bellii Western Painted Turtle X Turtles Emydidae Chrysemys picta dorsalis Southern Painted Turtle X Turtles Emydidae Deirochelys reticularia chrysea Florida Chicken Turtle X Turtles Emydidae Glyptemys insculpta Wood Turtle N Turtles Emydidae Graptemys barbouri Barbour's Map Turtle X Turtles Emydidae Graptemys ouachitensis Ouachita Map Turtle X N Turtles Emydidae Graptemys pseudogeographica False Map Turtle X Turtles Emydidae Graptemys pseudogeographica kohnii Mississippi Map Turtle X X

Table 6. All nonindigenous amphibian and reptile introductions into U.S. Fish and Wildlife Service Region 4 by state/territory. (Continued) Group Family Name Common Name AL AR FL GA KY LA MS NC PR SC TN VI Turtles Emydidae Trachemys dorbigni Brazilian Slider X Turtles Emydidae Trachemys scripta Pond Slider X Turtles Emydidae Trachemys scripta callirostris Columbian slider X Turtles Emydidae Trachemys scripta elegans Red-eared Slider X X N X Turtles Emydidae Trachemys scripta scripta Yellow-bellied Slider X Turtles Emydidae Trachemys stejnegeri malonei Inagua Slider X Turtles Kinosternidae Kinosternon scorpioides Scorpion Mud Turtle X Turtles Kinosternidae Staurotypus salvinii * Pacific Coast Giant Musk Turtle N Turtles Kinosternidae Sternotherus carinatus Razor-backed Must Turtle X Turtles Pelomedusidae Pelusios subniger East African Black Mud Turtle X Turtles Pelomedusidae Podocnemis lewyana Magdalena River Turtle X Turtles Pelomedusidae Podocnemis sextuberculata Six-tubercled River Turtle X Turtles Pelomedusidae Podocnemis unifilis Yellow-spotted River Turtle X Turtles Trionychidae Apalone ferox Florida Softshell X X N Turtles Trionychidae Apalone spinifera Spiny Softshell N

Bold = New to U.S. Fish and Wildlife Service Region 4 since 2000 * = New to United States X = Present in or prior to 2000 N = New since 2000

Species Accounts of Amphibians and Reptiles New to Region 4

Bombina orientalis (Boulenger, 1890) (Oriental Fire-bellied Toad)

Habitat: Bombina orientalis is semi-aquatic. It spends most of its time on land. The species usually lives in mixed coniferous/broad-leaved forests; however, it often occurs in other habitats, including spruce, pine or leafed forests, open meadows, river valleys, swampy bushlands, etc. Bombina orientalis inhabits different types of bodies of water with stagnant (sometimes running) water: lakes, ponds, swamps, streams, springs, ditches, puddles, etc. At the end of summer, the species can be found on land at distances up to few hundred Wikimedia meters from water (Kuzmin and others 2004; AmphibiaWeb, 2012).

Life History: This toad hibernates from late September/October to late April/May, usually on land in rotten trees, heaps of stones, leaves, and in groups of 1–6 individuals. Sometimes hibernation occurs in streams. Reproduction occurs from May to mid-August. The clutch contains 38–257 eggs deposited in portions of 3–45 eggs that the female attaches to plants near the water’s edge. The interval between depositing subsequent portions is 7–10 days. Females probably are capable of depositing multiple clutches. Embryonic and larval development takes about 2 months, and hatching takes place usually from early June to late July. Tadpoles complete metamorphosis usually by the end of August/late September. Maximum longevity is estimated as 20 years (Harkewicz, 2004; AmphibiaWeb, 2012).

Larvae consume detritus, various algae, fungi, higher plants, protozoans, and, in smaller amounts, aquatic invertebrates. Preying upon terrestrial invertebrates starts before the completion of metamorphosis. Adult food consists of terrestrial invertebrates including worms, mollusks, and insects. The proportion of aquatic invertebrates consumed varies among populations (Harkewicz, 2004; AmphibiaWeb, 2012).

Some predatory birds and mammals are known as natural enemies of this species. The defensive posture of adult individuals is similar to that of the European Fire-bellied Toad (B. bombina) (AmphibiaWeb, 2012).

Native Range: Korea, northeastern China and adjacent parts of Russia (Krysko and others, 2011).

Nonindigenous Range: Seven individuals have been captured from September 2007 to January 2010 at a single location in Broward County, Florida (UF 152327, 157280) (Krysko and others, 2011); however, Krysko, and others (2011) reported there is no evidence of reproduction.

Impacts: Unknown.

Comments: The bright, ventral colors serve as a warning to predators of toxicity. When the frog is disturbed or frightened, the toxin is secreted through the skin, on the back, hind legs, and sometimes the belly, in a milk-like substance. Not only will they emit this toxin, they also will lie on their back to show the color of the belly (the unkenreflex), indicating its toxicity to any predators (Harkewicz, 2004).

This species commonly is sold in large pet store chains. Because of the increased interest in outdoor water gardens, it may be at high risk of being introduced. Reportedly, this species can be kept outside in temperate regions of the U.S. (Harkewicz, 2004), implying that it can survive.

Afrixalus fornasini (Bianconi, 1849) (Fornasini’s Spiny Reed Frog)

Habitat: Afrixalus fornasinii lives in the savanna, possibly preferring dense savanna with larger bushes and trees. It is found in the coastal lowlands, but can be found up to 1,300 m. To the south, it is confined to coastal tropical lowlands of South Africa. Moist savannah, bush land, grassland, and dry forest are inhabited. Breeding occurs in permanent and semi- permanent pools, and marshes with reeds and sedges. (Pickersgill, and others, 2004; AmphibiaWeb, 2012).

Life History: Breeding begins in late September, and large choruses have been recorded in December and January. During the breeding season, A. fornasinii inhabits stagnant water bodies containing large stands of saw grass (Cyperus immensus) and cattails (Typha latifolia). Males may call Phil Bishop from elevated positions on sturdy, broad-leaved vegetation or from

44 reeds; calling begins shortly after sunset and continues until around midnight. The call consists of an initial vibrating pulse followed by a rapid stutter. It has been compared to the sound of a small machine gun (Harper, and others, 2010; Encyclopedia of Life, 2012a).

Eggs are laid over permanent water on reeds that are folded and glued over the eggs. Clutches contain approximately 80 unpigmented eggs. Hatching occurs 5 to 10 days later. The tadpoles are long and slender with flattened heads and a total length reaching 65 mm. The larval stage can last 75 days (Encyclopedia of Life, 2012a).

Native Range: Southeastern Africa, Kenya, Malawi, Mozambique, South Africa, Tanzania, and the United Republic of Zimbabwe (Krysko and others, 2011).

Nonindigenous Range: A single Foransini’s Spiny Reed Frog was captured in Broward County, Florida, in May 2010 (UF 163085) (Krysko and others, 2011).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown

Vector: Pet trade.

Impacts: Unknown. It has been reported that A. fornasini preys on eggs and developing larvae of other frogs (Drewes and Altig, 1996).

Comments: The skin of A. fornasini contains chemicals called tachykinins, which deter some mammalian predators (Channing and Howell, 2006).

Kaloula pulchra Gray, 1831 (Malaysian Painted Frog or Chubby Frog)

Habitat: This species is found in wetlands, river banks, forest edges, and dry forests, as well as agricultural and residential areas. It breeds in season pools or ponds (Kuangyang and others, 2009).

Alfeus Liman 45

Life History: The Malaysian Painted Frog is a nocturnal, fossorial species. It can burrow or climb into shrubby trees (Kuangyang and others, 2009).

In India, this frog calls after the first heavy monsoon showers in April through May. The males call while floating in pools of water. The tadpoles can metamorphose in as little as 2 weeks. This species eats worms and insects such as crickets, grasshoppers, and ants (AmphibiaWeb, 2012).

Native Range: Southeast Asia, Bangladesh, Cambodia, China, Hong Kong, India, Indonesia, Lao People's Democratic Republic, Malaysia, Myanmar, Nepal, Singapore, Thailand, and Vietnam (AmphibiaWeb, 2012).

Nonindigenous Range: A single individual was collected from Broward County, Florida, in January 2011 (UF 164746) (Krysko and others, 2011).

Impacts: Unknown. The species is a potential invasive species. It has been introduced and become established in Taiwan, Guam, Singapore, Borneo, and Sulawesi (formerly Celebes), with specimens noted in Australia and New Zealand (Lazell and Lu, 1996; Christy and others, 2007; Kuangyang and others, 2009).

Comments: It has been accidentally introduced to several countries (e.g., Guam), presumably via maritime or air-transport vessels and the pet trade (Christy and others, 2007). One specimen was discovered at the Perth airport in 2005, and one found in cargo at a New Zealand wharf, but there are no records showing the species has become established in either Australia or New Zealand. It often takes advantage of urban environments where it can be quite abundant (AmphibiaWeb, 2012).

The Malaysian Painted Frog can exude highly sticky secretions that most likely are used to deter predators. These secretions are noxious (unpleasant tasting) but do not contain detectable levels of toxins when tested (AmphibiaWeb, 2012).

Pipa pipa (Linnaeus, 1758) (Suriname Toad)

Habitat: Completely aquatic, found in sluggish rivers and canals with muddy bottoms (AmphibiaWeb, 2012).

Life History: This frog is entirely aquatic. Females lay approximately 100 eggs. After the male fertilizes the eggs, he attaches them to the female’s back and her skin grows around the eggs. Direct development (meaning they hatch as miniature frogs without going through a tadpole stage) occurs while on the M. Teixeira Jr. female’s back and the young emerge after 3 to 4 months (Rabb and Snedigar 1960; Rabb and Rabb, 1963).

Pipa pipa is omnivorous and eats worms, insects, crustaceans, and small fishes. It possessed long fingers, which are used to search for food. Once found, the fingers are used to stuff the food into its mouth (Wandzel, 1999).

Native Range: South America: Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Suriname, Trinidad and Tobago, and Venezuela (AmphibiaWeb, 2012).

Nonindigenous Range: A single individual was collected from a creek in Celada Ward, eastern Puerto Rico in August 2007.

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown

Vector: Pet release.

Impacts: Unknown.

Amphiuma tridactylum Cuvier, 1827 (Three-toed Amphiuma)

Habitat: The Three-toed Amphiuma often is found in bottomland marshes and lakes, bayous, cypress sloughs, and streams in hilly regions, frequently occupying crayfish burrows (Conant, 1975; Behler and King, 1979; Conant and Collins, 1998; Petranka, 1998). It is especially abundant in drainage ditches in suburban and agricultural areas of the lower Mississippi River (Behler and King, 1979). This amphiuma will move overland during and following heavy rains (Behler and King, 1979; Conant and Collins, 1998). It Matthew Niemiller spends most of the time hiding in heavily vegetated areas of permanent bodies of slow moving water, such as swamps, ponds, and lakes.

Life History: The adult Three-toed Amphiuma is capable of burrowing through soft mud and utilizing crayfish burrows. It can remain in these refugia for many months without feeding during droughts. This species often moves on land during nights with heavy rainstorms. They have been observed 12 m from the water’s edge (Petranka, 1998).

Amphiuma tridactylum is a nocturnal carnivore. Crawfish constitute a large proportion of the diet, followed by annelids, insects, mollusks, fishes, small turtles, and sometimes skinks along the shoreline (Chaney 1951; Fontenot and Fontenot, 1989). This species is preyed upon by Mudsnakes (Faracina abacura) and Cottonmouths (Agkistrodon piscivorous) (Behler, 1979; Petranka, 1998).

Mating occurs from December to June, and nesting from April to October. Unlike other amphiumas, this species has internal fertilization (Kreeger, 1942; Cagle, 1948; Petranka, 1998). The females are capable of storing viable sperm for at least 6–8 months (Kreeger 1942). Clutch size ranges from 44–282 with an average of 201 (Fontenot, 1999). Eggs are deposited in the water near land in coiled strings and are guarded by the female until hatching. The eggs generally are exposed to air as the water recedes. The larval stage lasts a mere 3 weeks. Sexual maturity is reached in 3–4 years (Petranka, 1998).

Native Range: The distribution of Amphiuma tridactylum is within the Gulf Coast Plain, centered on the lower Mississippi River from Texas to western Alabama, and north to southern Illinois and extreme southwestern Kentucky (Behler, 1979).

Nonindigenous Range: Several Three-toed Amphiumas were seen either seen or collected from a single location in Broward County, Florida, in December 2009 (UF 157220, 157286, 163094) (Krysko and others, 2011).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: Pet trade. Several dozen per year are sold from Louisiana as pets, whereas several hundred to about 1,000 are sold annually to biological suppliers for laboratory experiments. Most are taken as off- catch during crayfish trapping.

Impacts: Unknown.

Comments: Amphiumas are known to inflict a painful bite and also are known to flee when capture is attempted (Dundee and Rossman, 1989; Petranka, 1998). They also are covered with mucus slime and have thick skin, making them extremely difficult to grasp. There are two native species of amphiuma in Florida, the one-toed (A. pholeter) and the two-toed (A. means). As the common names imply, they can be distinguished by the number of toes on each leg.

Cynops orientalis (Boie, 1826) (Oriental Fire-bellied Newt)

Habitat: Found in low- and middle-elevation foothills ranging from 30 to 1,000 m above sea level, associated with bodies of still water. The species inhabits still ponds, rice terraces, or ditches in forest and degraded habitats. The substrate often is muddy. Flourishing aquatic vegetation makes good habitat for hiding and oviposition (Fei and others, 2006 in AmphibiaWeb, 2012).

Life History: In Zhengjiang Province, China, where the Henk Wallays winter is mild, this newt does not hibernate and can be found all year around. In other populations, individuals stay under water or in crevices on the ground over winter (Fei and others, 2006 in AmphibiaWeb, 2012).

The breeding season begins in March and ends in July, with a peak from late April to middle May. The suitable water temperature is 15–23 °C (Fei and others, 2006 in AmphibiaWeb, 2012). The female lays a single egg (or 2–4 eggs) on the ventral surface of aquatic leaf 2–10 centimeters below the water. She may fold the leaf with her legs to conceal the egg. Most eggs are laid during night. A single female can

49 lay around 100 eggs in 1 breeding season. The eggs hatch in 13–24 days. Males and females leave the breeding water from July to September. The annual male to female ratio is 2.3:1 (Fei and others, 2006 in AmphibiaWeb, 2012; Yang and Shen 1993 in AmphibiaWeb, 2012; Encyclopedia of Life, 2012a).

Cynops orientalis feeds on small aquatic animals such as worms and insect larvae (Fei and others, 2006 in AmphibiaWeb, 2012).

Native Range: Cynops orientalis is distributed widely in China in the lower reach of the Yangtze River and adjacent area, including Henan, Hubei, Anhui, Jiangsu, Zhejiang, Jiangxi, and Fujian Provinces (Fei and others, 2006 in AmphibiaWeb, 2012).

Nonindigenous Range: Four collections of single individuals in Broward County, Florida: one on June 4, 2009; the second on August 22, 2009; the third on June 21, 2010; and the fourth on July 3, 2010 (UF 157033, UF 157946, UF 158946). A fifth specimen was collected in Sumter County, Florida, in July 2010 (UF 158927) (Krysko and others, 2011).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: Pet trade. Oriental Fire-bellied Newts are one of the most commonly found species in the pet trade due to extensive collection from the wild, where hundreds of thousands of individuals are collected annually. Cynops orientalis commonly is used either for educational purposes or as research material (Fei and others, 2006 in AmphibiaWeb, 2012).

Impacts: Unknown. Oriental Fire-bellied Newts are mildly poisonous and excrete toxins through their skin. Consisting primarily of tetrodotoxins, newts of the genus Cynops pose a medically significant threat if enough toxins are consumed (Brodie and others, 1974).

Comments: The animals are sold by the thousands in pet markets in China and Europe. This species is used for educational and research purposes; in traditional Chinese medicine it is used to cure skin itching or burning (Fei and others, 2006 in AmphibiaWeb, 2012).

Pachytriton labiatus (Unterstein, 1930) (Paddle-tail Newt)

Habitat: The Paddle-tail Newt inhabits low-gradient streams of varying sizes. It breeds in the same habitats and the larvae develop in the water. The water can be either clear or murky as a result of flooding with a substrate of either sand or small rocks (Zhao and others 1994 in AmphibianWeb 2012).

Life History: The species is fully aquatic. Populations in Hunan Province, China, hibernate from November till April in crevices or under boulders where water is Hou Mian present. The newts hide in shelters during daytime and are active at night. Pachytriton labiatus feeds on earthworms, other aquatic arthropods, and insect larvae. Anti-predator postures include stretching of limbs, lifting the head and tail, and exposing the orange ventral blotches (Zhao and others 1994 in AmphibiaWeb, 2012; Sparreboom and Wu, 2013). The breeding season typically begins in April and lasts to July. Females lay around 40 single eggs at the lower surface of rocks in streams; sometimes the eggs are clustered to form a patch (Fei et al, 2006 in AmphibiaWeb, 2012; Sparreboom and Wu, 2013).

Native Range: Central and southern China (Krysko and others, 2011).

Nonindigenous Range: One specimen was collected from Broward County, Florida, in February 2010 (UF 157219) (Krysko and others, 2011).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: Pet trade. Large numbers of wild-caught animals appear in the pet trade every year. Local inhabitants may sell them as juvenile Giant Chinese Salamanders (Andrias davidianus) (Xu and others 2002 in AmphibiaWeb, 2012).

Impacts: Unknown.

Comments: Paddletail Newts sometimes are confused with the Japanese Fire-bellied Newts (C. pyrrhogaster) and sold as such, but are much larger, more aggressive, and will eat smaller Fire-bellies if given the opportunity (Encyclopedia of Life, 2012c).

Paleosuchus palpebrosus (Cuvier, 1807) (Cuvier's Dwarf Caiman)

Habitat: This species can be found near rivers and inundated savanna areas. It has been known to occupy streams of varying sizes, where it is spotted resting near the shorelines. This species is also terrestrial, and can be seen on piles of small rocks and, river banks, and near decaying trees. P. palpebrosus is known to dwell in burrows up to 1.5- 3.5 meters long. This species prefers colder temperatures and can survive as low as 6 °C (Stevenson, 1999). Topí Pigula Life History: Paleosuchus palpebrosus primarily feeds on fishes as an adult, although they also take insects, frogs, crustaceans, and rodents. There is little data available on the breeding habits of this species. In Colombia, they nest between August and November during the rainy season; however, it appears that nesting occurs in the dry season in Brazil. Their mound-like nests have been found on floating vegetation, but most often are on the forest floor. The female lays between 13 and 15 eggs. Incubation is 90–92 days long (Stevenson, 1999).

Native Range: Cuvier's Dwarf Caiman is native to South America. It is most commonly found in Brazil, French Guiana, Surinam, Guyana, and Venezuela (Grenard, 1991).

Nonindigenous Range: A single specimen was collected from Long Pine Key in Everglades National Park, Miami-Dade County, Florida, in August 2008 (EVER 40566; UF 153469) (Krysko and others, 2011).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: Pet trade.

Impacts: Unknown.

Crocodylus acutus (Cuvier, 1807) (American Crocodile)

Habitat: In Florida, C. acutus prefers brackish waters and coastal mangrove swamps but may wander inland or out to sea (Ashton and Ashton, 1991; Moler, 1992). One population of American Crocodiles exists in the canal system of a Florida power plant (Gaby and others, 1985; Grenard, 1991). Adults are fairly tolerant of high salinity levels, but the young may be dependent on drinking from lenses of freshwater that temporarily float on heavier saltwater following rainfall (Moler, 1992). Thomas Castelazo Life History: The diet consists mostly of crabs, fish, snakes, turtles, birds, and small mammals (including dogs), with insects and spiders included in the diet of juveniles (Moler, 1992). In Florida, mating occurs in late winter and early spring (Moler, 1992). Females lay hard-shelled eggs in April or May, which are buried in a simple hole or mound of soil or on a beach, stream bank, or canal levee (Gaby and others, 1985; Kushlan and Mazzotti, 1989b; Grenard, 1991; Moler, 1992). Females are not as diligent about protecting the nest as Alligator mississippiensis, but will open the nest at hatching and transport young to the water in their mouths (Kushlan and Mazzotti, 1989b; Moler, 1992).

Native Range: The American crocodile ranges from the Atlantic and Pacific coasts of Mexico; Central American to northern South America and the Caribbean islands of Cuba, Jamaica, Hispaniola; and the southern tip of Florida (Thorbjarnarson, 1989). In North America, C. acutus lives at the northernmost extent of its range, found only along the extreme southern coastline of Florida, including the Keys, primarily confined to the counties of Dade and Monroe, with individuals wandering as far north as Palm Beach County on the east coast and Sarasota County on the west coast (Kushlan and Mazzotti, 1989a).

Nonindigenous Range: A single juvenile was collected at Tortuguero Lagoon, Puerto Rico, in May 2004.

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown

Vector: Pet release. Although the island of Puerto Rico is outside the normal native range for this species, an adult could theoretically have swum to Puerto Rico; however, because the individual

53 collected was a juvenile, it is believed to be the result of a pet release. This same area also hosts a population of spectacled caiman (C. crocodylus) that resulted from pet releases (Lever, 2003; Kraus,, 2009).

Impacts: Unknown.

Comments: Although adult American Crocodiles potentially are dangerous to humans, they are not prone to unprovoked attacks like some other species of the genus Crocodylus (Moler, 1992; Conant and Collins, 1998). Crocodylus acutus is protected under the U.S. Endangered Species Act, both federally and in the State of Florida (Moler, 1992; Mitchell, 1994; Levell, 1997).

Eunectes murinus (Linnaeus, 1758) (Green Anaconda)

Habitat: The Green Anaconda is an aquatic snake usually found submerged under the water or near the water’s edge. It can be found in deep, shallow, turbid, and clear waters, and lacustrine and riverine habitats. It often is found in seasonally flooded areas (Reed and Rodda, 2009). Because of its aquatic nature, it is able to escape extremes in temperature variation.

Life History: This species is a constrictor, nonvenomous, and an ambush predator. It immobilizes and kills its prey by LA Dawson using its jaws to hold on the prey while coiling its body around the animal and constricting until it dies from suffocation or other crushing injuries. The list of potential prey items of an anaconda is extensive and varied, and consists of birds, amphibians, reptiles, mammals, and fish. Small individuals may climb trees to raid bird nests. The Green Anaconda swallows its prey whole, even prey much larger than the diameter of their mouths. They are known to consume large prey such as peccaries, capybaras, tapirs, deer, and sheep. This species has a slow-acting digestive system; it often takes days or weeks to digest food. After a meal, an anaconda may not eat for weeks or months (Reed and Rodda, 2009). There are few records of attacks on humans, and no well-documented deaths (Rivas, 1998; Reed and Rodda, 2009).

Anacondas are not immune from predators. Neonates and juveniles are particularly prone to predation, and seem to suffer high mortality within their first year. Predators of small anacondas include Crab- eating Foxes, tegu lizards, Crested Caracaras, caimans, and larger Green Anacondas (Rivas and others, 1999; Rivas and others, 2001; Mattison, 2006).

The Green Anaconda reproduces sexually and has internal fertilization. Courtship often extends over several months and the mating period typically is from April to May. The female is thought to lay down a

54 pheromone trail, which attracts the male. Often the snakes cluster in a breeding ball which may consist of up to 13 males coiled around one female. The gestation period for the Green Anaconda is 6 to 10 months. This species gives birth to live young. A female anaconda can give birth to as many as a 82 young, though typically the size of the litter ranges from 28 to 42 (Reed and Rodda, 2009).

Native Range: Green Anacondas are found throughout tropical South America, east of the Andes, mainly in the Amazon and Orinoco basins and in the Guianas, as well as the island of Trinidad (Reed and Rodda, 2009).

Nonindigenous Range: In August 2004, a Green Anaconda (UF 143935) was collected from Big Cypress Swamp, Collier County, Florida. A juvenile male was collected in December 2004 from Fakahatchee Strand Preserve State Park in Collier County. An adult was capture in January 2010 at East Lake Fish Camp in Osceola County (Krysko and others, 2011). Another individual was collected in Osceola Park, Florida, in August 2010. More anecdotal reports exist for the Everglades area. There is the possibility of an established population in the Fakahatchee Strand area (Krysko and others, 2011).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: Pet trade.

Impacts: Unknown.

Comments: Adults of this species can grow to over 25 feet in length. For its length, it is much heavier than other large constrictors; therefore, if weight and length are both considered, Green Anacondas are the largest snakes in the world (Reed and Rodda, 2009).

Eunectes notaeus (Cope, 1862) (Yellow Anaconda)

Habitat: The Yellow Anaconda lives mainly in water and appears to be restricted to swampy, seasonally flooded, or riverine habitats (Reed and Rodda, 2009)

Life History: The Yellow Anaconda is a generalist carnivore, preying mainly on animals found in wetland and riparian areas throughout its range. Its diet consists of birds, bird eggs, small mammals,

55 Patrick Jean turtles, lizards, other snakes, occasional fish or fish carrion, capybara, and caimans (Reed and Rodda, 2009). The Yellow Anaconda is considered an ambush hunter and constrictor. The digestive system is relatively slow and the Yellow Anaconda may eat only every few days or months, depending on the size of the last prey item. The Yellow Anaconda can survive long periods without prey (Reed and Rodda, 2009)).

An adult Yellow Anaconda has few natural predators. Humans are the main predator and it is hunted primarily for its skin, although there are varying numbers imported each year for the pet trade. Predators of juveniles and the occasional adult include Crab-eating Foxes (Cerdocyon thous), tegu lizards (Tupinambis merianae), Spectacled Caimans (Caiman crocodilus), larger anacondas (Eunectes), felids (cats), canids (dogs), procyonids (raccoon family), mustelids (weasel/skunk family), herons, and raptors such as the Crested Caracara (Polyborus plancus) (Reed and Rodda, 2009).

The Yellow Anaconda is a solitary animal, except in breeding season (April and May). Yellow Anacondas have been known to form breeding balls, consisting of one female and multiple males. After a 6-month gestation period, the females give birth to fully developed live young. There is great variation in the literature about litter size, but Reed and Rodda (2009) believed the best estimate for wild individuals was 7–42 with an average of 19.5. These young immediately are able to live on their own. Young anacondas reach sexual maturity at 17 to 29 months old. In captivity, Yellow Anacondas have lived to more than 20 years of age (Reed and Rodda, 2009).

Native Range: The Yellow Anaconda occurs in southern South America, including the Pantanal in Bolivia and Brazil, and south through the Paraguay and Parana River Basins in Paraguay and Argentina (Reed and Rodda, 2009).

Nonindigenous Range: Two specimens collected from the same area in Wapanocca National Wildlife Refuge, Arkansas: one in the fall of 2004, the other in the spring of 2005. This could indicate that they were released together and one overwintered. Four specimens have been collected from Florida. One from the Tamiami Trail in the Everglades in 2006, two in Big Cypress National Preserve in 2007, and another in Big Cypress in early 2008. There are anecdotes of others collected near the Palm Beach County line (Reed and Rodda, 2009; Krysko and others, 2011).

Impacts: Unknown.

Erpeton tentaculatus Lacepede, 1800 (Tentacled Snake)

Habitat: This species inhabits stagnant or slow moving bodies of water with emergent-submergent vegetation (Uetz, 2012). During the dry season it will bury itself under the mud until the rainy season returns. It can be found in fresh, brackish, and sea water (Murphy and others, 2010).

Life History: This rear-fanged aquatic snake feeds exclusively on fish, which is accomplished via an ambush method. It lives its entire life in murky water and can stay underwater for up to 30 minutes without coming up for air. It can move Ryan Somma only awkwardly on land. In dry times and at night, the snake may burrow itself in the mud. The 5-13 young develop ovoviviparously (eggs hatch internally) and are born live underwater (Thiesen, 2012).

Native Range: This species is endemic to Peninsular Southeast Asia, occurring in Thailand and southern Vietnam (Uetz, 2012)

Nonindigenous Range: A single individual was collected near an importer’s facility in Hollywood, Florida, in December 2010 (UF 163086). No others were found and a breeding population is not suspected (Holbrook and Krysko, 2011; Krysko and others, 2011).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: Pet trade.

Impacts: Unknown.

Comments: Although it does have venomous fangs, the Tentacled Snake is not considered dangerous to humans (Thieson, 2012). The fangs are small, only partially grooved, and positioned deep in the rear of the mouth. This is the only species of snake to possess twin "tentacles" on the front of its head, which have been shown to have mechano-sensory function (Toronto Zoo, 2012).

Rhinoclemmys pulcherrima (Gray, 1855) (Painted Wood Turtle)

Habitat: The Painted Wood Turtle is found in moist woodlands and scrub often close to streams or ponds. The species also occurs in cleared or developed areas close to streams (Ernst and Barbour, 1989; Ernst and others, 2012).

Life History: Rhinoclemmys pulcherrima consumes wildflowers, grasses, fruit, insects, worms, and fish (Ernst and Barbour, 1989; Jolman, 2000). It nests in leaf litter or soil and lays several clutches of three to five eggs during May to December Tornadohalt (Ernst and others, 2012).

Native Range: The Painted Wood Turtle is found along the Pacific coast from Sonora, Mexico, to Costa Rica (Ernst and Barbour, 1989; Ernst and others, 2012).

Nonindigenous Range: One individual was captured in Manatee County, Florida, in August 2006 (UF 121604).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: Pet trade.

Impacts: Unknown.

Chelydra serpentina (Linnaeus, 1758) (Snapping Turtle)

Habitat: Chelydra serpentina is an adaptable, highly aquatic turtle that inhabits almost any body of water (including brackish), rarely basks, but will foray over land (Babcock, 1919; Gibbons and Semlitsch, 1991; Ernst and others, 1994).

Life History: The Snapping Turtle is a generalized omnivore; any invertebrate, vertebrate, aquatic plant, or carrion they can grab and swallow, will be consumed (Ernst and Barbour,

1989; Ernst and others, 1994). In northern regions it has a Gerald Sneegas high tolerance to cold and occasionally can be seen crawling beneath ice in winter (Ernst and others, 1994). Female C. serpentina lay 6–104 eggs in soil, rotting vegetation, sawdust piles, or muskrat and beaver lodges, during spring or summer (Ernst and others, 1994).

Native Range: The Snapping Turtle ranges from Nova Scotia, New Brunswick, and southern Quebec to southeastern Alberta, and southward east of the Rocky Mountains to Florida and Texas (Ernst and others, 1994).

Nonindigenous Range: Two adults were collected from Espiritu Santo River Natural Reserve in Puerto Rico in March 2006.

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown

Vector: Pet release.

Impacts: Unknown.

Comments: The Snapping Turtle derives its common name from its violent antipredator behavior, which involves snapping at an adversary while quickly lunging forward and extending its neck, potentially delivering a painful, damaging wound (Ernst and others, 1994; Oldfield and Moriarty, 1994; Hammerson, 1999). This vigorous defense is exhibited out of water and does not seem to be used while submerged (Vogt, 1981).

Macrochelys temminickii (Troost in Harlan, 1835) (Alligator Snapping Turtle)

Habitat: The Alligator Snapping Turtle is a massive, highly aquatic turtle that inhabits deep waters, primarily deep rivers with steep banks, but also lakes and swamps (Pritchard, 1989, 1992).

Life History: For the most part, M. temminckii is a carnivore that feeds primarily on fish, smaller turtles (especially kinosternids), crayfish, and mollusks (mussels and snails), but occasionally eats juvenile alligators, small mammals, ducks, amphibians, carrion, U.S. Fish and Wildlife Service fruit, and acorns (Dundee and Rossman, 1989; Ernst and Barbour, 1989; Pritchard, 1989; Ernst and others, 1994; Conant and Collins, 1998). Although they actively forage for food along the bottom, they hunt for fish by lying still in ambush, with open mouth while wriggling the lure appendage on the tongue (Obst, 1986, 1998; Pritchard, 1989, 1992; Zug and others, 2001). Fish are trapped in their mouths in this fashion more commonly by smaller, younger turtles than by larger adults that spend more time actively foraging (Pritchard, 1989). Females typically lay their eggs in loose, moist soil in or near river banks (Pritchard, 1989).

Native Range: This species is confined to river systems that drain into the Gulf of Mexico. It is found in almost all river systems from the Suwannee River, Florida, to eastern Texas and extends as far north as Kansas, Illinois, and Indiana (Ernst and others, 1994).

Nonindigenous Range: A single individual was collected in 2007 from Espiritu Santa Natural Reserve in Puerto Rico.

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown

Vector: Pet release.

Impacts: Unknown.

Glyptemys insculpta (LeConte, 1830) (Wood Turtle)

Habitat: The wood turtle inhabits wetlands, streams, and rivers, as well as meadows, forests, and farmlands adjacent to water (Ernst and others, 1994; Conant and Collins, 1998).

Life History: The species hibernates in water, and is most aquatic in the northern portion of its range (Ernst and others, 1994; Conant and Collins, 1998). Mating peaks in spring and fall and nesting occurs in May and June, when 4 to 18 eggs are laid in holes excavated in Phil Myers the soil (Ernst and others, 1994).

Native Range: The wood turtle is found in the northeastern United States, the Great Lakes region, New Brunswick, and Nova Scotia with isolated populations in northern New York and Quebec (Conant and Collins, 1998).

Nonindigenous Range: A single specimens was captured in St. Augustine, Florida, in August 2009.

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Vector: Pet release. Unknown

Impacts: Unknown.

Staurotypus salvinii Gray, 1864 (Pacific Coast Giant Musk Turtle)

Habitat: The Pacific Coast Giant Musk Turtle inhabits slow- flowing waters with soft bottoms and aquatic vegetation (Ernst and others, 2012).

Life History: Like other musk turtle species, they are carnivorous, eating various types of aquatic invertebrates, as

61 Dawson well as fish, amphibians, and carrion. Clutches consist of 6–10 eggs that incubate from 80–210 days (Ernst and others, 2012).

Native Range: Mexico (lowland Pacific drainages of Oaxaca and Chiapas), Guatemala, El Salvador, and Belize (Uetz, 2012).

Nonindigenous Range: Two individuals were collected in 2010 at Zoo Miami, Florida (UF 160342, UF 162276). The first was a gravid female in October; the next was a juvenile that was collected 1 month later less than a mile away (Smith and others, 2011).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: Pet trade.

Impacts: Unknown.

Apalone spinifera (Le Sueur, 1827) (Spiny Softshell)

Habitat: The Spiny Softshell inhabits small marshy creeks and farm ponds, and large, fast-flowing rivers and lakes. This species apparently requires a soft bottom and some aquatic vegetation, as well as sand bars or bands for basking (Behler and King, 1979; Ernst and others, 1994).

Life History: This species nests from May to August in a bank of sand or gravel exposed to full sun andlays 4-32 eggs that hatch in late August to October or overwinter in the nest until James Parham spring (Behler and King, 1979). Females produce two clutches per year (Ernst and others, 1994). Sex is not determined by temperature (Greenbaum and Carr, 2001). The Spiny softshell is carnivorous, preying on crayfish, fish, insects, carrion and other aquatic invertebrates (Ernst and others, 1994).

Native Range: This species is found in eastern and central North America, from portions of Ontario and New York, west through the plains states, south to the Gulf of Mexico and the southern states, through

62 limited sections of northern Florida, with disjunct populations in Montana, Wyoming, and other western states (Ernst and Lovich, 2009).

Nonindigenous Range: At least two individuals, an adult male and adult female, have been seen over a period of 3 years (2008–2010) at a pond on Florida International University, Miami-Dade County, Florida (Krysko and others, 2011).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: Pet trade.

Impacts: Unknown.

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FISH

Overview

There were 19 new fish species detected in the region since 2000 (Table 7 and Table 8), more than any other taxonomic group (Figure 1). Nineteen additional fish species introductions are the result of marine aquarium releases. Because the U.S. Fish and Wildlife Service does not manage marine species, these are mentioned, but freshwater species are the focus of this report. All marine aquarium fish were collected in Florida. The lionfish also was reported from all other coastal states in the Region. These species will not be discussed further; however, it should be noted that aquarium release is a major pathway and U.S. Fish and Wildlife Service does have jurisdiction over importation and inspection of shipments. The new freshwater fishes to the region and the first year and state where they were found are listed in Table 7. For the sake of brevity, only new species are listed on this table, unlike the other groups.

3 An average of 1.67 new fishes is introduced into the Region every year.

In every year except 2011, between one and 2 three new fish were introduced to the region (Figure 15).

2011 2012 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010

Figure 15. Number of new fish species to the region by year.

Puerto Rico and Florida were the two major entry points of new fish species to the region since 2001 (Figure 16).

Figure 16. Number of new fish species to the region and state of first occurrence.

Aquarium release is the most dominant 1 2 Aquarium release vector for new fish introductions (Figure 17). 1 Released bait

Escaped aquaculture Released food fish 15

Figure 17. Number of new introduced fish species by vector.

Nearly one-half of the new fish species introduced to the region come from South and Central America. Another quarter of the 2 South and Central species are from Asia and Eurasia (Figure America 18). North America 9 6 Asia/Eurasia

Africa

Figure 18. Number of new introduced fish species by continental origin.

There were 63 new state introductions of fishes. More new freshwater fish were reported from Puerto Rico (21) than any other territory or state in the Region (Figure 19). This largely is because a systematic island-wide survey took place in the mid-2000s that revealed many species that were not previously known from the Island. There was an average of 5.25 new species per state over the 12 year period, and new state records per year during 2001-2012.

Puerto Rico the territory that had the most new fish species introduced (Figure 19).

Figure 19. Number of new state/territory records per state/territory 2001-2012. At least four new fish species were introduced in 7 of the 12 years (Figure 20).

0 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 Figure 20. Number of new state/territory records per year.

Thirteen species were not only new to Region 4, but also new introductions to the country (Table 7).

Table 7. New nonindigenous fish species introductions into U.S. Fish and Wildlife Service Region 4 since 2000. First Continent of Population Group Family Common Name Year Origin Likely Vector Status Alestiidae Phenacogrammus interrupts * Congo Tetra 2006 Africa released aquarium established Channidae Channa maculata Blotched Snakehead 2009 Asia released food fish unknown Channidae Channa micropeltes Giant Snakehead 2005 Asia released aquarium failed Cichlidae Cichlasoma labiatum Red Devil 2006 South America released aquarium established Cichlidae Parachromis dovii* Wolf Cichlid 2009 Central America released aquarium established Cichlidae Paraneetroplus melanurus x P. zonatus * hybrid cichlid 2001 South America released aquarium unknown Cichlidae Paraneetroplus synspilus Redhead Cichlid 2010 Central America released aquarium established Cyprinidae Mylopharyngodon piceus Black Carp 2004 Asia escaped captivity aquaculture unknown White Cloud Mountain Cyprinidae Tanichthys albonubes * Minnow 2008 Asia released aquarium unknown Gasterosteidae Apeltes quadracus Fourspine Stickleback 2002 North America released bait unknown Gyrinocheilidae Gyrinocheilus aymonieri* Chinese Algae-eater 2007 Asia released aquarium established Lepisosteidae Atractosteus spatula Alligator Gar 2010 North America released aquarium unknown Loricariidae Ancistrus sp. * Bristlenosed Catfish 2001 South America released aquarium established Loricariidae Farlowella vittata* Stick Catfish 2012 South America released aquarium unknown Loricariidae Glyptoperichthys gibbiceps * Leopard Pleco 2007 South America released aquarium unknown Loricariidae Pterygoplichthys anisitsi * Southern Sailfin Catfish 2003 South America released aquarium established Osteoglossidae Osteoglossum bicirrhosum Arawana 2003 South America released aquarium failed Polypteridae Polypterus delhezi * Barred Bichir 2001 Africa released aquarium failed Tetraodontidae Tetraodon nigroviridis * Spotted Green Pufferfish 2004 Asia escaped captivity aquaculture failed * = New to United States

Table 8. All nonindigenous fish species introductions into U.S. Fish and Wildlife Service Region 4 by state/territory. Family Name Common Name AL AR FL GA KY LA MS NC PR SC TN VI Acipenseridae Acipenser transmontanus White Sturgeon X Alestiidae Phenacogrammus interruptus * Congo Tetra N Amiidae Amia calva Bowfin X X Anabantidae Anabas testudineus Climbing Perch X Anabantidae Ctenopoma nigropannosum Twospot Climbing Perch X Anostomidae Leporinus fasciatus Banded Leporinus X Belontiidae Trichopsis vittata Croaking Gourami X Callichthyidae Callichthys callichthys Cascarudo X Callichthyidae Corydoras sp. Corydoras X Callichthyidae Hoplosternum littorale Brown Hoplo X Catostomidae Carpiodes cyprinus Quillback X Catostomidae Carpiodes velifer Highfin Carpsucker X X Catostomidae Catostomus commersonii White Sucker X X Catostomidae Hypentelium etowanum Alabama Hog Sucker X Catostomidae Hypentelium nigricans † Northern Hog Sucker X X Catostomidae Ictiobus bubalus Smallmouth Buffalo X X Catostomidae Ictiobus cyprinellus Bigmouth Buffalo X X N Catostomidae Moxostoma lachneri Greater jumprock X Catostomidae Moxostoma rupiscartes Striped Jumprock X X Centrarchidae Ambloplites cavifrons Roanoke Bass X Centrarchidae Ambloplites constellatus Ozark Bass X Centrarchidae Ambloplites rupestris Rock Bass X X X Centrarchidae Enneacanthus gloriosus Bluespotted Sunfish X Centrarchidae Lepomis auritus Redbreast Sunfish X X X X X X X X Centrarchidae Lepomis cyanellus Green Sunfish X X X X X Centrarchidae Lepomis gibbosus Pumpkinseed X X X X Centrarchidae Lepomis gulosus Warmouth X X X Centrarchidae Lepomis humilis Orangespotted Sunfish X X X

Table 8. All nonindigenous fish species introductions into U.S. Fish and Wildlife Service Region 4 by state/territory. (Continued)

Family Name Common Name AL AR FL GA KY LA MS NC PR SC TN VI Centrarchidae Lepomis macrochirus Bluegill X X X X Centrarchidae Lepomis megalotis Longear Sunfish X Centrarchidae Lepomis microlophus Redear Sunfish X X X X X Centrarchidae Lepomis miniatus Redspotted Sunfish X Centrarchidae Micropterus cataractae Shoal Bass X Centrarchidae Micropterus coosae Redeye Bass X X X X X X Centrarchidae Micropterus dolomieu Smallmouth Bass X X X X X X X Centrarchidae Micropterus punctulatus Spotted Bass X X X X X X X Centrarchidae Micropterus punctulatus henshalli † Alabama Spotted Bass X X Centrarchidae Micropterus punctulatus punctulatus † Northern Spotted Bass X X X Centrarchidae Micropterus salmoides Largemouth Bass X X X Centrarchidae Micropterus salmoides floridanus † Florida Largemouth Bass X X Centrarchidae Pomoxis annularis White Crappie X X X X X X Centrarchidae Pomoxis nigromaculatus Black Crappie X X X Channidae Channa argus † Northern Snakehead N X N Channidae Channa maculata Blotched Snakehead N Channidae Channa marulius † Bullseye Snakehead X Channidae Channa micropeltes Giant Snakehead N Characidae Aphyocharax anisitsi Bloodfin Tetra X Characidae Astyanax mexicanus Mexican Tetra X Characidae Colossoma macropomum Tambaqui X N X Characidae Colossoma or Piaractus sp. unidentified pacu N X X X X X X Characidae Gymnocorymbus ternetzi Black Tetra X N Characidae Hyphessobrycon eques Serpae Tetra X Characidae Metynnis sp. Metynnis X X Characidae Moenkhausia sanctaefilomenae Redeye Tetra X Characidae Piaractus brachypomus Pirapatinga, Red-bellied Pacu X X X X X N X X N N Characidae Piaractus mesopotamicus Small-scaled Pacu X Characidae Pygocentrus nattereri Red Piranha X

Table 8. All nonindigenous fish species introductions into U.S. Fish and Wildlife Service Region 4 by state/territory. (Continued)

Family Name Common Name AL AR FL GA KY LA MS NC PR SC TN VI Characidae Pygocentrus or Serrasalmus sp. unidentified piranha X X Characidae Serrasalmus rhombeus White Piranha X Cichlidae Aequidens pulcher Blue Acara X Cichlidae Amphilophus citrinellus Midas Cichlid X N Cichlidae Astronotus ocellatus Oscar N X X X X N X N Cichlidae Cichla ocellaris Butterfly Peacock Bass X X Cichlidae Cichla temensis Speckled Pavon X Cichlidae Cichlasoma bimaculatum Black Acara X Cichlidae Cichlasoma citrinellum x C. urophthalmus * hybrid cichlid N Cichlidae Cichlasoma labiatum Red Devil N Cichlidae Cichlasoma nigrofasciatum Convict Cichlid N X N N Cichlidae Cichlasoma octofasciata Jack Dempsey X Cichlidae Cichlasoma salvini Yellowbelly Cichlid X Cichlidae Cichlasoma trimaculatum Threespot Cichlid X Cichlidae Cichlasoma urophthalmus Mayan Cichlid X Cichlidae Geophagus sp. Eartheater X Cichlidae Haplochromis callipterus † Eastern Happy X Cichlidae Hemichromis letourneuxi African Jewelfish X N Cichlidae Herichthys cyanoguttatum † Rio Grande Cichlid X X N Cichlidae Heros severus Banded Cichlid X Cichlidae Labeotrpheus sp. scrapermouth cichlid X Cichlidae Oreochromis aureus Blue Tilapia X X X X X X X Cichlidae Oreochromis mossambicus Mozambique Tilapia X X X X X N Cichlidae Oreochromis mossambicus x O. hornorum † hybrid tilapia X Cichlidae Oreochromis niloticus Nile Tilapia X X X X N X Cichlidae Oreochromis urolepis Wami Tilapia X Cichlidae Oreochromis urolepis hornorum x O. mossambicus † hybrid tilapia X Cichlidae Oreochromis, Sarotherodon, Tilapia sp. tilapia X X X X X Cichlidae Oreochromis, Sarotherodon, Tilapia sp. x sp. hybrid tilapia X

Table 8. All nonindigenous fish species introductions into U.S. Fish and Wildlife Service Region 4 by state/territory. (Continued)

Family Name Common Name AL AR FL GA KY LA MS NC PR SC TN VI Cichlidae Parachromis dovii * Wolf Cichlid N Cichlidae Parachromis managuensis Jaguar Guapote X N N Cichlidae Paraneetroplus melanurus x P. zonatus * hybrid cichlid N Cichlidae Paraneetroplus synspilus * Redhead Cichlid N Cichlidae Pterophyllum scalare Freshwater Angelfish X X Cichlidae Pterophyllum sp. freshwater angelfish X Cichlidae Sarotherodon melanotheron Blackchin Tilapia X Cichlidae Telmatochromis bifrenatus Lake Tanganyika Dwarf Cichlid X Cichlidae Thorichthys meeki Firemouth Cichlid X N Cichlidae Tilapia buttikoferi † Zebra Tilapia X Cichlidae Tilapia mariae Spotted Tilapia X Cichlidae Tilapia rendalli Redbreast Tilapia X Cichlidae Tilapia sparrmanii Banded Tilapia X Cichlidae Tilapia zillii Redbelly Tilapia X X X X X Clariidae Clarias batrachus Walking Catfish X X Cobitidae Misgurnus anguillicaudatus Oriental Weatherfish N X N N X Cobitidae Pangio kuhlii Coolie Loach X Cottidae Cottus bairdii Mottled Sculpin X Cyprinidae Barbonymus schwanenfeldii Tinfoil Barb X Cyprinidae Brachydanio rerio Zebra Danio X Cyprinidae Campostoma anomalum Central Stoneroller X X Cyprinidae Campostoma oligolepis Largescale Stoneroller X Cyprinidae Carassius auratus Goldfish X X X X X X X X X X X Cyprinidae Clinostomus funduloides funduloides † Rosyside Dace X Cyprinidae Ctenopharyngodon idella Grass Carp X X X X X X X X X X X Cyprinidae Ctenopharyngodon idella x Hypophthalmichthys nobilis † Grass Carp x Bighead Carp X Cyprinidae Cyprinella galactura Whitetail Shiner X X N Cyprinidae Cyprinella lutrensis Red Shiner X X X X Cyprinidae Cyprinella pyrrhomelas Fieryblack Shiner X

Table 8. All nonindigenous fish species introductions into U.S. Fish and Wildlife Service Region 4 by state/territory. (Continued)

Family Name Common Name AL AR FL GA KY LA MS NC PR SC TN VI Cyprinidae Cyprinella venusta Blacktail Shiner X Cyprinidae Cyprinella venusta cercostigma † Eastern Blacktail Shiner X Cyprinidae Cyprinus carpio Common Carp X X X X X X X X N X X Cyprinidae Devario malabaricus Malabar Danio X Cyprinidae Hybognathus hankinsoni Brassy Minnow X Cyprinidae Hybopsis winchelli Clear Chub X Cyprinidae Hypophthalmichthys molitrix Silver Carp X X X X X X X Cyprinidae Hypophthalmichthys molitrix x H. nobilis † hybrid silver/bighead carp X X Cyprinidae Hypophthalmichthys nobilis Bighead Carp X X X X X X N X Cyprinidae Labeo chrysophekadion Black Sharkminnow, Black Labeo X Cyprinidae Leuciscus idus Ide X X X X X X Cyprinidae Luxilus albeolus White Shiner X Cyprinidae Luxilus cerasinus Crescent Shiner X Cyprinidae Luxilus chrysocephalus isolepis Striped Shiner X X Cyprinidae Luxilus coccogenis Warpaint Shiner X X Cyprinidae Luxilus zonistius Bandfin Shiner X X Cyprinidae Lythrurus atrapiculus Blacktip Shiner X X Cyprinidae Lythrurus fasciolaris † Scarlet Shiner X Cyprinidae Lythrurus matutinus † Pinewoods Shiner X Cyprinidae Mylopharyngodon piceus Black Carp N N Cyprinidae Nocomis biguttatus Hornyhead Chub X Cyprinidae Nocomis leptocephalus Bluehead Chub X X X Cyprinidae Nocomis micropogon River Chub X X X Cyprinidae Nocomis raneyi Bull Chub X Cyprinidae Notemigonus crysoleucas Golden Shiner X X X X X Cyprinidae Notropis amoenus Comely Shiner X Cyprinidae Notropis baileyi Rough Shiner X X X Cyprinidae Notropis bifrenatus Bridle Shiner X Cyprinidae Notropis buccatus Silverjaw Minnow X

Table 8. All nonindigenous fish species introductions into U.S. Fish and Wildlife Service Region 4 by state/territory. (Continued)

Family Name Common Name AL AR FL GA KY LA MS NC PR SC TN VI Cyprinidae Notropis chiliticus Redlip Shiner X N Cyprinidae Notropis chrosomus Rainbow Shiner X X Cyprinidae Notropis harperi Redeye Chub X Cyprinidae Notropis hudsonius Spottail Shiner X Cyprinidae Notropis hypsilepis Highscale Shiner X Cyprinidae Notropis leuciodus Tennessee Shiner X X X Cyprinidae Notropis longirostris Longnose Shiner X Cyprinidae Notropis lutipinnis Yellowfin Shiner X X Cyprinidae Notropis ozarcanus Ozark Shiner X Cyprinidae Notropis potteri Chub Shiner X X Cyprinidae Notropis procne Swallowtail Shiner X Cyprinidae Notropis rubricroceus Saffron Shiner X Cyprinidae Notropis shumardi Silverband Shiner X Cyprinidae Notropis spectrunculus Mirror Shiner X X Cyprinidae Notropis telescopus Telescope Shiner X X Cyprinidae Notropis texanus Weed Shiner X X X Cyprinidae Notropis xaenocephalus Coosa Shiner X Cyprinidae Phoxinus oreas Mountain Redbelly Dace X X X Cyprinidae Pimephales notatus Bluntnose Minnow X Cyprinidae Pimephales promelas Fathead Minnow X X X X X X X X X X X Cyprinidae Puntius conchonius Rosy Barb X N X Cyprinidae Puntius gelius Dwarf Barb X Cyprinidae Puntius tetrazona Tiger Barb X N Cyprinidae Rhinichthys atratulus Blacknose Dace X Cyprinidae Rhinichthys cataractae Longnose Dace X Cyprinidae Scardinius erythrophthalmus Rudd X X Cyprinidae Tanichthys albonubes * White Cloud Mountain Minnow N Cyprinidae Tinca tinca Tench X X X X X X X X X X Cyprinodontidae Jordanella floridae Flagfish X

Table 8. All nonindigenous fish species introductions into U.S. Fish and Wildlife Service Region 4 by state/territory. (Continued)

Family Name Common Name AL AR FL GA KY LA MS NC PR SC TN VI Doradidae Oxydoras niger Ripsaw Catfish X Doradidae Platydoras costatus Raphael Catfish X Doradidae Pterodoras granulosus Granulated Catfish X Doradidae Pterodoras sp. Thorny Catfish X Erythrinidae Hoplias malabaricus Trahira X Esocidae Esox lucius Northern Pike X X X X X Esocidae Esox lucius x E. masquinongy † Tiger Muskellunge X X Esocidae Esox masquinongy Muskellunge X X X X X X X Esocidae Esox niger Chain Pickerel X X X Fundulidae Fundulus catenatus Northern Studfish X X Fundulidae Fundulus lineolatus Lined Topminnow X Fundulidae Fundulus seminolis Seminole Killifish X X Fundulidae Fundulus stellifer Southern Studfish X Fundulidae Lucania goodei Bluefin Killifish X X Gadidae Lota lota † Burbot X Gasterosteidae Apeltes quadracus Fourspine Stickleback N Gasterosteidae Culaea inconstans Brook Stickleback X X X Gyrinocheilidae Gyrinocheilus aymonieri * Chinese Algae-eater N Helostomatidae Helostoma temminkii Kissing Gourami X Heptapteridae Rhamdia quelen Bagre X Ictaluridae Ameiurus brunneus Snail Bullhead X X Ictaluridae Ameiurus catus White Catfish X X X X X X X Ictaluridae Ameiurus melas Black Bullhead X X N X Ictaluridae Ameiurus nebulosus Brown Bullhead X X X X X Ictaluridae Ameiurus platycephalus Flat Bullhead X X X Ictaluridae Ictalurus furcatus Blue Catfish X X X X X X Ictaluridae Ictalurus punctatus Channel Catfish X X X X X X Ictaluridae Noturus exilis Slender Madtom X Ictaluridae Noturus insignis Margined Madtom X X

Table 8. All nonindigenous fish species introductions into U.S. Fish and Wildlife Service Region 4 by state/territory. (Continued)

Family Name Common Name AL AR FL GA KY LA MS NC PR SC TN VI Ictaluridae Pylodictis olivaris Flathead Catfish X X X X Lepisosteidae Atractosteus spatula Alligator Gar N Loricariidae Ancistrus sp. * Bristlenosed Catfish N Loricariidae Farlowella sp. * Stick Catfish N Loricariidae Glyptoperichthys gibbiceps * Leopard Pleco N Loricariidae Hypostomus plecostomus Plecostomus X N Loricariidae Hypostomus sp. suckermouth catfish X X Loricariidae Pterygoplichthys anisitsi * Southern Sailfin Catfish N Loricariidae Pterygoplichthys disjunctivus Vermiculated Sailfin Catfish X X N X Loricariidae Pterygoplichthys multiradiatus Orinoco Sailfin Catfish X N Loricariidae Pterygoplichthys pardalis Amazon Sailfin Catfish X X X Loricariidae Pterygoplichthys sp. sailfin catfish X Mastacembelidae Macrognathus siamensis † Spotfin Spiny Eel X Moronidae Morone americana x M. saxatilis † White Perch x Striped Bass X Moronidae Morone mississippiensis Yellow Bass X X X X Moronidae Morone mississippiensis x M. saxatilis † Yellow Bass x Striped Bass X Notopteridae Chitala ornata Clown Knife X N Osphronemidae Betta splendens Siamese Fighting Fish X Osphronemidae Colisa fasciata Banded Gourami X Osphronemidae Colisa labiosa Thicklipped Gourami X Osphronemidae Colisa lalia Dwarf Gourami X Osphronemidae Macropodus opercularis Paradise Fish X X Osphronemidae Osphronemus goramy Giant Gourami X Osphronemidae Trichogaster leerii Pearl Gourami X Osphronemidae Trichogaster trichopterus Three-spot Gourami X N Osteoglossidae Osteoglossum bicirrhosum Arawana N Pangasiidae Pangasius hypophthalmus † Iridescent Shark X Percidae Ammocrypta bifascia Florida Sand Darter X Percidae Etheostoma edwini Brown Darter X

Table 8. All nonindigenous fish species introductions into U.S. Fish and Wildlife Service Region 4 by state/territory. (Continued)

Family Name Common Name AL AR FL GA KY LA MS NC PR SC TN VI Percidae Etheostoma fusiforme Swamp Darter X Percidae Etheostoma nuchale Watercress Darter X Percidae Etheostoma olmstedi Tessellated Darter X Percidae Etheostoma zonale Banded Darter X Percidae Perca flavescens Yellow Perch X X X X X X X X X Percidae Percina maculata † Blackside Darter X X Percidae Percina nigrofasciata † Blackbanded Darter X X Percidae Percina tanasi Snail Darter X Percidae Sander canadensis Sauger X X X X X X X X Percidae Sander canadensis x S. vitreus † Saugeye X X X Percidae Sander vitreus Walleye X X X X X X X X X Pimelodidae Phractocephalus hemioliopterus Redtail Catfish X N Poeciliidae Belonesox belizanus Pike Killifish X Poeciliidae Gambusia affinis Western Mosquitofish X X X X X Poeciliidae Gambusia holbrooki Eastern Mosquitofish X X X X X Poeciliidae Heterandria formosa Least Killifish X Poeciliidae Poecilia hybrid Lyretail Black Molly X Poeciliidae Poecilia latipinna x P. velifera Black Molly X Poeciliidae Poecilia latipunctata Tamesi Molly X Poeciliidae Poecilia petenensis Peten Molly X Poeciliidae Poecilia reticulata Guppy X X N Poeciliidae Poecilia sphenops Mexican Molly X N Poeciliidae Poecilia vivipara † Guaru X Poeciliidae Xiphophorus hellerii Green Swordtail X N X Poeciliidae Xiphophorus hellerii x X. maculatus † Red Swordtail X Poeciliidae Xiphophorus hellerii x X. variatus † platyfish/swordtail X PoeciliidaeTable 8. AllXiphophorus nonindigenous maculatus fish species introductions into U.S.S Fishouthern and Platyfish Wildlife Service Region 4. (Continued) X N X Poeciliidae Xiphophorus maculatus x X. variatus † platyfish/swordtail X

Poeciliidae Xiphophorus sp. † Platyfish X

Table 8. All nonindigenous fish species introductions into U.S. Fish and Wildlife Service Region 4 by state/territory. (Continued)

Family Name Common Name AL AR FL GA KY LA MS NC PR SC TN VI Poeciliidae Xiphophorus variatus Variable Platyfish X Poeciliidae Xiphophorus xiphidium † Swordtail Platy X Polyodontidae Polyodon spathula Paddlefish X X Polypteridae Polypterus delhezi * Barred Bichir N Salmonidae Coregonus artedi Cisco X Salmonidae Oncorhynchus clarkii Cutthroat Trout X X Salmonidae Salmo letnica Ohrid Trout X Salmonidae Salmo trutta x Salvelinus fontinalis Tiger Trout N X Salmonidae Salvelinus fontinalis Brook Trout X X X X X X Salmonidae Salvelinus namaycush Lake Trout X X X Schilbeidae Platytropius siamensis False Siamese Shark X Synbranchidae Monopterus albus Asian Swamp Eel X X Tetraodontidae Tetraodon nigroviridis * Spotted Green Pufferfish N Bold = New to U.S. Fish and Wildlife Service Region 4 since 2000 * = New to United States † = Introduced to USFWS Region 4 prior to 2001 but not included in last report X = Present in or before 2000 N = New since 2000

The seven fish species that were found in more than one new state or territory, indicating they are spreading or have been introduced multiple times are listed in Table 9. Five of these species (Oscar, Convict Cichlid, Oriental Weatherfish, Jaguar Guapote, and Red-bellied Pacu) are in the aquarium trade and likely are releases by individuals. Three of the five (Oscar, Jaguar Guapote, and Red-bellied Pacu) grow to large sizes, which likely is the reason for their release. The fourth, Convict Cichlid, is aggressive towards tank mates. Northern Snakehead likely is a result of an illegal food fish release in North Carolina and of an aquaculture escape in Arkansas. Black Carp also are aquaculture escapes and have become established in the Mississippi River.

Table 9. Fish species found as “new” in more than one state or territory. Genus Species Common Name State/Terr Year Astronotus ocellatus Oscar AR 2005 NC 2011 SC 2004 Channa argus Northern Snakehead AR 2008 NC 2002 Cichlasoma nigrofasciatum Convict Cichlid AL 2011 LA 2004 PR 2006 Misgurnus anguillicaudatus Oriental Weatherfish AL 2009 LA 2005 NC 2009 Mylopharyngodon piceus Black Carp AR 2005 LA 2004 Parachromis [=Cichlasoma] managuense Jaguar Guapote LA 2004 PR 2007 Piaractus brachypomus Red-bellied Pacu LA 2002 SC 2003 TN 2010 Although different species of Sailfin Catfish (Pterygoplichthys spp. and others) have been found in new states and territories, functionally they are a single group. These fish, also known in the aquarium trade as “plecos,” are purchased by aquarium keepers to eat algae from aquarium surfaces, including the glass. They are not actively marketed as different species and even most fish taxonomists cannot distinguish them from one another (Table 10).

Table 10. Sailfin catfish species sold as algae eaters to aquarists. Genus Species Common Name State/Terr Year Pterygoplichthys anisitsi Southern Sailfin Catfish FL 2003 Pterygoplichthys disjunctivus Vermiculated Sailfin Catfish NC 2002 Pterygoplichthys multiradiatus Orinoco Sailfin Catfish PR 2007 Glyptoperichthys gibbiceps Leopard Pleco PR 2007 Hypostomus plecostomus Suckermouth Catfish PR 2004

Species Accounts of Fishes New to Region 4

Phenacogrammus interruptus (Boulenger, 1899) (Congo Tetra)

Habitat: The Congo Tetra is benthopelagic and is found in waters with a pH between 6.0 and 8.0 (Froese and Pauly, 2012).

Life History: The Congo Tetra feeds on worms, small insects, crustaceans, and plant matter. The female lays up to 300 eggs, which sink to the bottom and hatch in 6 days. The nest is not guarded (Mills and Vevers, 1989). André Karwath

Males are larger and more colorful than females (SeriouslyFish, 2012a).

Native Range: This species is native to the Middle Congo Basin in Africa (Froese and Pauly, 2012).

Nonindigenous Range: The Congo Tetra is established in Naranjito Creek (Turabo River), Puerto Rico, as of 2006. This species is not listed by Erdman (1984) as present in Puerto Rico.

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown

Vector: Aquarium release.

Impacts: Unknown.

Channa maculata (Lacepède, 1801) (Blotched Snakehead)

Habitat: The Blotched Snakehead lives in streams, lakes, ponds, and ditches and prefers shallow water with vegetation (Courtenay and Williams, 2004).

Life History: This species builds circular, open nests in vegetation where the eggs float to the surface and are guarded by the parents. It feeds on crustaceans, large insects, frogs, and fishes using ambush tactics (Courtenay and Williams, 2004). Chinese Academy of Fishery Sciences Native Range: This snakehead is found in Asia, including Middle Japan, southern China, Vietnam, Taiwan, and the Philippines (Froese and Pauly, 2012).

Nonindigenous Range: A specimen originally misidentified as Channa argus was collected from Lake Wylie, North Carolina, in 2009 (NCSM 53258; W. Starnes, personal communication).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: The most likely mechanism is as a released food fish, because it is common in the food trade (Courtenay and Williams, 2004). It is rarely available in the aquarium trade (Courtenay and Williams, 2004).

Impacts: Unknown. This predatory fish has the potential to impact native fauna. This species has been introduced to Madagascar where it is considered to contribute to the decline of amphibian populations (Courtenay and Williams, 2004).

Channa micropeltes (Cuvier in Cuvier and Valenciennes, 1831) (Giant Snakehead)

Habitat: The Giant Snakehead prefers lakes, reservoirs, canals, and rivers—most commonly deep, standing or slow flowing water (Courtenay and Williams, 2004).

Life History: It nests in a circular area that the parents clear of vegetation. The eggs rise in the water column where they are guarded by parents even after they hatch. The Giant Snakehead is a daytime predator, feeding on fishes, frogs, Thesupermat and birds. It has a reputation of being very aggressive and even attacking humans (Courtenay and Williams, 2004).

Native Range: This species of snakehead is from Southeast Asia including India, possibly Burma (now Myanmar), Thailand, Malay Peninsula, Sumatra, Borneo, Java, and Billiton (Roberts, 1989), as well as Laos (Baird and others, 1999).

Nonindigenous Range: A single fish collected from Poplar Tree Lake, Tennessee, in December 2005, was originally misidentified as C. argus.

Impacts: Unknown.

Comments: This species would likely establish in Florida if introduced (Courtenay and Williams, 2004).

Amphilophus labiatus (Günther, 1864) (Red Devil)

Habitat: The Red Devil inhabits lakes and rarely enters rivers (Conkel, 1993).

Atelier Monpli 86

Life History: It feeds on small fish, snails, insect larvae, worms, and other bottom-dwelling organisms (Yamamoto and Tagawa, 2000). The female tends to the clutch of eggs (Froese and Pauly, 2012).

Native Range: The Red Devil is native to tropical America, including theAtlantic Slope drainages in Central America, including Great Lakes of Nicaragua (Conkel 1993).

Nonindigenous Range: In Puerto Rico, Red Devils are first collected in Puerto Rico in 2006 and are established in the Loiza, Dos Bocas and Caonillas Reservoirs.

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown

Vector: Aquarium release.

Impact: This species has been reported to have “some adverse ecological effects” and “some undecided socio-economic effect” (Froese and Pauly, 2012).

Parachromis dovii (Günther, 1864) (Wolf Cichlid)

Habitat: The Wolf Cichlid inhabits lakes but also thrives in various lower and middle river valleys. It prefers tropical temperatures of 21–37 °C (Froese and Pauly, 2012).

Life History: This species is an avid cavern digger. It is piscivorous, although it also eats crustaceans and insects in smaller numbers. It is considered to have medium fecundity with a minimum population doubling time of 1.4–4.4 years (Froese and Pauly, 2012).

AfricanCichlidsFish.blogspot.com

Native Range: The Wolf Cichlid is native to Central America including the Atlantic slope, from the Aguan River (Honduras) to the Moín River (Costa Rica); and the Pacific slope, from the Yeguare River (Honduras) to the Bebedero River (Costa Rica) (Froese and Pauly, 2012).

Nonindigenous Range: Established in Loiza Reservoir in eastern Puerto Rico since at least 2009.

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown

Vector: Aquarium release.

Impacts: Unknown.

Paraneetroplus melanurus x Paraneetroplus zonatus (hybrid cichlid)

Habitat: Paraneetroplus melanurus inhabits sandy-bottomed lakes (Froese and Pauly, 2012). No specific information was found for P. zonatus.

Life History: It feeds on aufwuchs, benthic detritus of vegetable matter, and small crustaceans. It is a nest guarder that produces 300–500 fry per spawning (Froese and Pauly, 2012). No information was available for P. zonatus.

Native Range: P. melanurus is native to Central America: Atlantic slope, in the De la Pasión and Lake Petén Basins, Guatemala. P. zonatus is native to Central America: Pacific slope, Tehuantepec River basin, Mexico (Froese and Pauly, 2012).

Nonindigenous Range: Hybrid cichlids identified as 'Theraps (now Paraneetroplus) Hybrids were collected in Biscayne Canal, Miami-Dade County, Florida, between 2001 and 2006 (Shafland and others, 2008a, b).

Impacts: Unknown.

Comments: Historically, cichlid taxonomy has been problematic due to a proliferation of species and genera with few uniquely identifying characteristics, and movement or reassignment of taxa with little justification. Due to copius research, cichlid taxonomy, systematics, and nomenclature have changed rapidly over the past decade. Many species formerly placed in the genus Theraps (= Vieja), including T. (V.) melanurus and T. (V.) zonatus, were moved by McMahan and others (2010) to Paraneetroplus based on molecular data.

Paraneetroplus synspilus (Hubbs, 1935) (Redhead Cichlid)

Habitat: The Redhead Cichlid inhabits lakes and the lower river valley with a slight tolerance for the brackish environment (Froese and Pauly, 2012).

Life History: This species is found in tropical waters with temperatures ranging from 24–30 °C. It produces a maximum of 1,000 eggs, and sexual maturity is reached at 10 cm (Froese and

Pauly, 2012). Frank Teigler

Native Range: The Redhead Cichlid is native to Central America, including the Atlantic slope, in the Usumacinta River drainage in Mexico, Guatemala, and Belize (Froese and Pauly, 2012).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown

Nonindigenous Range: Established in Guajataca Reservoir in northern Puerto Rico since at least 2012.

Vector: Aquarium release.

Impacts: Unknown.

Mylopharyngodon piceus (Richardson, 1846) (Black Carp)

Habitat: This species can be found in rivers, streams, or lakes; however, it requires large rivers to reproduce (Nico, 2005).

Life History: Reproduction takes place in late spring and summer when water temperatures and/or water levels rise (Nico and others, 2005). Both male and female black carp are broadcast spawners; females are capable of releasing hundreds of thousands of eggs into flowing water, which then develop in the pelagic zone Leo Nico (Nico and others, 2005). After fertilization, the eggs become semiboyant (Sukhanova, 1967 as cited in Nico and others, 2005). They hatch in 1 to 2 days, depending on water temperatures, and the yolk sac is absorbed in 6 to 8 days (Nico and others, 2005). They become sexually mature at 4 to 6 years after which they migrate back to their spawning grounds (Nico and others, 2005). Successful reproduction is known only from riverine habitats (Nico and others, 2005). Lifespan probably is greater than 15 years (Biro, 1999 as cited in Nico and others, 2005).

Native Range: The Black Carp is native to most major Pacific drainages of eastern Asia from the Pearl River (Zhu Jiang) Basin in China north to the Amur River (Heilong Jiang) Basin of China, and far eastern Russia; possibly native to the Honghe or Red River of northern Vietnam (Nico and others, 2005).

Nonindigenous Range: A single fish was collected from the White River, Arkansas, in April 2005 (Nico and others, 2005). Recent information indicates that commercial fishers have been collecting Black Carp on an annual basis since the early 1990s in the Red and Atchafalaya Rivers and the lower Mississippi River main channel in Louisiana (Nico and others, 2005). At least 12 of the captured specimens were later examined by biologists, but many others have been captured over the past 15 years (see section on status) (Nico and others, 2005; L. G. Nico, U.S. Geological Survey, unpub. Data, 2013). Recent documented captures in Louisiana consist of three specimens captured in May 2007; these include two fish taken from the Red-Atchafalaya systems and one from the main channel of the Mississippi River (L. G. Nico, unpub. data, 2013). Commercial fishers reported taking a Black Carp in early July 2004 from the Mississippi River near the mouth of the Homochitto River in Mississippi (Nico and others, 2005). Outside the southeast region of the United States, approximately 30 Black Carp were reported as having escaped from a fish farm in Missouri into the Osage River, Missouri River Basin, in April 1994 (Nico and others, 2005). At least three specimens have been documented from Illinois open waters. The first was captured on March 26, 2003, from Horseshoe Lake, Alexander County, in the southern part of the state (Chick and others, 2003). A second specimen was taken June 10, 2004, from the Mississippi River near Lock and Dam 24 near Clarksville Island side channel in Calhoun County (Nico and others, 2005). A third fish was found July 1, 2010, in Pool 25 of the Mississippi River near Hamburg, Illinois.

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: This species was first brought into the United States in the early 1970s as a "contaminant" in imported Grass Carp stocks. These fish came from Asia and were sent to a private fish farm in Arkansas (Nico and others, 2005). Subsequent introductions of Black Carp into this country occurred in the early 1980s. During this period it was imported as a food fish and as a biological control agent to combat the spread of Yellow Grub Clinostomum margaritum in aquaculture ponds (Nico and others, 2005). There is evidence that large portions of the lower Mississippi River Basin where aquaculture farms are present have been subject to large-scale floods on a number of occasions over the past few decades. Consequently, it is likely that these farm escapes are the source of some or all of the Black Carp present in the lower Mississippi River Basin (Nico and others, 2005:245). There also is risk that Black Carp may be spread by other means. According to one aquaculture farmer, hundreds of young Black Carp accidentally were included in shipments of live baitfish sent from Arkansas to bait dealers in Missouri as early as

1994 (Nico and others, 2005:5). In addition, because of the continued widespread distribution of Grass Carp across the United States, the possibility remains that shipments may inadvertently contain Black Carp (Nico and others, 2005). Juveniles, in particular, are difficult to distinguish from Grass Carp young. As such, Nico and others (2005) expressed concern over the increased risk that the species be misidentified and unintentionally introduced as Grass Carp to some areas.

Impacts: There is high potential that the Black Carp would negatively impact native aquatic communities by feeding on, and reducing, populations of native mussels and snails, many of which are considered endangered or threatened (Nico and others, 2005). Black Carp have the potential to restructure benthic communities by direct predation and removal of algae-grazing snails. Mussel beds consisting of smaller individuals and juvenile recruits probably are most vulnerable to being consumed (Nico and others, 2005). Furthermore, based on the fact that Black Carp attain a large size (well over 1 meter long), both juvenile and adult mussels and snails of many species would be vulnerable to predation by this fish (Nico and others, 2005). Fish farmers report that Black Carp are very effective in reducing the numbers of snails in some ponds. Wui and Engle (2007) argued that Black Carp can eliminate 100 percent of the snails in a single pond. Although their assumption that Black Carp are capable of eliminating all common pond snails in ponds is open to debate, their effectiveness in significantly reducing snail populations in aquaculture ponds indicates that any Black Carp occurring in the wild may cause significant declines in certain native mollusk populations in North American streams and lakes (Nico and others, 2005). Because the life span of Black Carp reportedly is over 15 years, sterile triploid Black Carp in the wild would be expected to persist many years and therefore have the potential to cause harm to native mollusks by way of predation (Nico and others, 2005).

Comments: Chick and others (2003) believed that their Illinois capture was the first wild record of Black Carp in the United States, but Nico and others (2005) provided new information indicating that Louisiana commercial fishers had been collecting Black Carp in Louisiana since the early 1990s; however, until recently the Louisiana commercial fishers thought that the Black Carp in their nets were just an unusual type of Grass Carp—somewhat darker and with a higher dorsal fin and more pointed head or snout (Nico and others, 2005, p. xiii). The Black Carp that escaped in Missouri during the 1994 floods may have been triploid and thus considered sterile; however, it also was rumored that these fish may have been brood stock. All wild Black Carp examined to date taken in Louisiana waters have been found to be diploid (Nico and others, 2005).

The Black Carp is a bottom-dwelling molluscivore that has been used by U.S. fish farmers to prey on and control disease-carrying snails in their farm ponds; more recently, this species has been proposed as a biological control for the introduced Zebra Mussel Dreissena polymorpha. Although the subject has been debated, currently, there is no experimental evidence that indicates Black Carp would be effective in controlling Zebra Mussels. Because Black Carp do not have jaw teeth and their mouths are relatively small, it is unlikely that these fish are capable of breaking apart Zebra Mussel rafts (Nico and others, 2005).

Tanichthys albonubes (Lin, 1932) (White Cloud Mountain Minnow)

Habitat: This species inhabits clear, slow-moving brooks with thick weed growth (Chan and Chen, 2009).

Life History: The White Cloud Mountain minnow is cold water tolerant fish often kept in aquaria. Although the nominal temperature range for the species in the wild is 18–22 °C, it can survive water temperatures down to 5 °C (Froese and Pauly, 2012). It feeds on plankton, meiobenthos and larvae of aquatic insects. This species spawns Jan Ševčík several times a year, from March to October, on plants (Cui, 2011).

Native Range: First discovered at White Cloud Mountain (also known as Mount Baiyun or Baiyunshan) a few miles north of the city of Guangzhou, Guangdong Province, China and thought to be restricted to the surrounding Pearl River Delta region. Unfortunately it has probably been extirpated from that locality; the “mountain” is actually a collection of 30 or so peaks and is now a very popular tourist resort complete with cable car access, hotels, and public parks. Between 1980 and 2001, the species went unrecorded in nature, leading to fear of its extinction (Seriously Fish, 2012b).

Nonindigenous Range: This species was collected in a tributary of the Savannah River northwest of Augusta, Georgia, in 2008. No one has checked on the population since its discovery to know its status.

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: Aquarium release.

Impacts: Unknown.

Apeltes quadracus (Mitchill, 1815) (Fourspine Stickleback)

Habitat: The Fourspine Stickleback can be found in fresh and brackish water from 0 to 3 m deep. Adults inhabit weedy bays and backwaters (Delbeek and Williams, 1988). Dave Neely Life History: Apeltes quadracus often live sympatrically with other species of stickleback throughout their native range, though they generally are solitary with regard to other members of their own species. Individuals often are observed perched among bottom debris and vegetation, never in open water, hovering over a patch of substrate, picking at it frequently, then moving on to another patch (Delbeek and Williams, 1987). Delbeek and Williams (1987) also found that Fourspine Stickleback fed exclusively on the bottom, either among benthic vegetation or from the substrate, the most abundant organisms in their diets being diatoms, nauplii, nematodes, ostracods, and cyclopods. During spawning, the males of this species establish territories and build small nests in which females lay their eggs (Courtenay, 1985). The male then chases the female away and cares for the eggs until they hatch. Courtenay (1985) also found that, unlike other stickleback species, male Apeltes tend multiple nests at one time in laboratory and natural breeding site observations. The Fourspine Stickleback also has the widest range of salinity tolerance of any North American species of stickleback (Holm and Hamilton, 1988).

Native Range: Europe and Atlantic coast of North America. Atlantic Slope from Gulf of St. Lawrence to Trent River system, North Carolina. Mostly a near-shore marine species, but far inland populations occur in lakes in Nova Scotia; in Hudson, Delaware; and in Susquehanna River drainages (Page and Burr, 2011).

Nonindigenous Range: Found in Lake Jackson, Alabama, in 2002.

Impacts: Unknown in Alabama, but rapid increases of Apeltes in Thunder Bay suggest the species is displacing native stickleback species at a rapid rate (Stephenson and Momot, 2000).

Gyrinocheilus aymonieri (Tirant, 1883) (Chinese Algae-eater)

Habitat: The Chinese Algae-eater is found in medium- to large-sized rivers and flooded fields, and in mountain streams with gravel bottoms (Froese and Pauly, 2012).

Life History: This species is freshwater, demersal, and tropical, preferring temperatures of 25–28 °C. It mostly is herbivorous but also takes insect larvae or zooplankton. Its sucker-like mouth is used to hold onto fixed Kjell Nilsson objects in currents (Froese and Pauly, 2012).

Native Range: This species is native to Asia, including the Mekong, Chao Phraya, and Meklong Basins and the northern Malay Peninsula (Froese and Pauly, 2012).

Nonindigenous Range: Chinese Algae-eaters were first collected in Rio Canos, in eastern Puerto Rico, just upstream of Lago Loiza, in the spring of 2007 (Wayne Starnes, written communication, 2007).

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown

Vector: Aquarium release. These fish are very popular in the aquarium trade for controlling algae in the tank.

Impacts: Unknown.

Atractosteus spatula (Lacepède, 1803) (Alligator Gar)

Habitat: The Alligator Gar lives in coastal estuaries and major coastal rivers (Mettee and others, 1996). It is found in fresh, brackish, and occasionally salt water (Boschung and Mayden, 2004).

Konrad P. 95 Schmidt

Life History: The Alligator Gar is the largest of the gars, reaching nearly 10 feet in length (Ross, 2001). It feeds primarily on fishes, but will also eat crabs, carrion, and birds. This species spawns in late spring; the eggs are toxic (Ross, 2001).

Native Range: Mississippi River Basin from southwestern Ohio and southern Illinois south to Gulf of Mexico; Gulf Coastal Plain from Econfina River, Florida, to Veracruz, Mexico (Page and Burr, 1991).

Nonindigenous Range: A single Alligator Gar was collected in July 2010 in Lake Wateree, South Carolina.

Impacts: Unknown.

Ancistrus sp. Kner, 1854 (Bristlenosed Catfish) possibly A. cf. cirrhosus

Habitat: Species in this genus are found throughout rivers and floodplain areas. Ancistrus cf. cirrhosus inhabits streams ranging from still, turbid water with clay substrate to free-flowing, clear water with gravel substrate (Froese and Pauly, 2012)

Life History: The unique tentacles are thought to be used in sensing speed and direction of stream K. Jinkings currents and perhaps in detecting odors (Burgess, 1989). Another hypothesis is that they resemble juvenile fish and are used to attract females (Sabaj, and others, 1999). Only the males develop the bull, bushy tentacles (Sabaj, and others, 1999).

This species has an algae diet, typical for Loricariid catfishes (Froese and Pauly, 2012).

Ancistrus species have the capability of obtaining oxygen by breathing air their modified stomach. This allows them to survive in conditions with low oxygen levels (Gee 1976; Sabaj, and others, 1999).

Breeding takes place in hollows, caves, and mud holes in banks. The female may lay 20–200 adhesive eggs, usually to the ceiling of the cavity. The male takes care of the young. During this time, a male usually will not leave the cavity to feed, or will leave occasionally and quickly return. The eggs hatch in 4–10 days; the male guards the eggs for 7–10 days after hatching. The fry remain in the cave and become free swimming in 2–4 days (Sabaj, and others, 1999).

Native Range: Species of this genus are native to the Amazon River in South America (Froese and Pauly, 2012). There are 64 described, valid species of Ancistrus (Froese and Pauly, 2012). The species A. cf. cirrhosus is native to the Argentinian portion of the Parana River drainage (Froese and Pauly, 2012).

Nonindigenous Range: An unidentified species of Ancistrus has been collected from Tamiami Canal, Florida, from 2001–2005, and from Snapper Creek Canal in 2006 (Shafland and others, 2008a, b).

Impacts: Unknown.

Comments: According to aquarium information, it appears that the species found is most likely A. cf. cirrhosus. This is the most common species in the trade; however, no one really knows the origin of the stock or even if they are still pure. They may be a hybrid. This species so commonly is raised in captivity that it is called Ancistrus sp. 3 (SeriouslyFish, 2012c).

Farlowella vittata Myers, 1942 (Stick catfish)

Habitat: Stick catfish are found in slow moving rivers and streams (Covain and Fisch-Muller, 2007). Aquarium information suggests keeping this species in a dimly lit, well-oxygenated stream set-up with a sand substrate, rocks, and lots of branches. The species prefers temperatures of 75–80 °F (SeriouslyFish, 2012d).

Life History: Stick catfishes, along with most other loricariid catfishes, feed primarily on algae and detritus. Species of Farlowella primarily Wikimedia Commons inhabit slow moving rivers and streams, camoflaged among submerged sticks and leaves (Covain and Fisch-Muller, 2007). Farlowella exhibits some degree of parental care in reproduction, with males guarding eggs deposited on open vertical surfaces including submerged vegetation or rocks (Burgess, 1989, Covain and Fisch-Muller, 2007). Eggs hatch in 6–10 days (SeriouslyFish, 2012d).

Native Range: South America. Farlowella vittata is found in the Orinoco River Basin in Colombia and Venezuela (Retzer and Page, 1997; Froese and Pauly, 2012).

Nonindigenous Range: A single fish was collected in the Hillsborough River, Florida, in April 2012.

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: Aquarium release.

Impacts: Unknown.

Comments: Farlowella vittata, along with F. acus and F. gracilis, commonly is found in the aquarium trade.

Glyptoperichthys gibbiceps (Kner, 1854) (Leopard Pleco)

Habitat: The Leopard Pleco can be found in tropical freshwaters from 23–27 °C (Froese and Pauly, 2012).

Life History: The Leopard Pleco is nocturnal. It feeds on algae (Ferraris 2007), although it will also feed on dead animals. Egg rearing is also presumed to take place in burrows dug into mud banks, similar to many related species. Soulkeeper Native Range: South America: Middle and upper Amazon and Orinoco basins (Froese and Pauly, 2012).

Nonindigenous Range: Collected from Dos Bocas Reservoir, Puerto Rico, in 2007.

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown

Vector: Aquarium release.

Impacts: Unknown, but probably similar to Pterygoplichthys spp. (a synonym of the genus).

Comment: As of 1989, this species was the second most imported Loricariid, after Pterygoplichthys anisitsi (Burgess, 1989). That probably is no longer the case.

Pterygoplichthys anisitsi Eigenmann and Kennedy, 1903 (Southern Sailfin Catfish)

Habitat: Pterygoplichthys spp. can be found in a wide variety of habitats, ranging from relatively cool, fast-flowing, and oxygen-rich highland streams to slow-flowing, warm lowland rivers and stagnant pools poor in oxygen. They are tropical fish and populations typically are limited only by their lower lethal temperature, which has been found to be about 8.8–11 °C in some species (Gestring, 2006). They can thrive in a range of acidic to alkaline waters in a pH range of about 5.5.0 to 8.0 (Mendoza and others, 2009). They often are found in soft waters, but can adapt very quickly to hard Bob Howells waters. Pterygoplichthys spp. also are highly tolerant to poor water quality and commonly are found in polluted waters (Chavez and others, 2006). They are known to use outflow from sewage treatment plants as thermal refugia and can readily adapt to changing water quality (Nico & Martin, 2001). Some species are salt tolerant (Mendoza and others, 2009).

Life History: Freshwater; demersal; Tropical; 21–24 °C (Froese and Pauly, 2012). This catfish primarily is a detritivore, but it also feed on benthic invertebrates and algae. Like other members of this group, it digs burrows for nests (Nico and Martin, 2001)

Native Range: South America: Paraguay, middle Paraná, Bermejo, and Uruguay River Basins (Froese and Pauly, 2012).

Nonindigenous Range: The Southern sailfin catfish has been collected in the St. Johns River Basin, Florida.

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown Vector: Aquarium release.

Impacts: Largely unknown. In several natural streams, this species is relatively abundant. Because it grazes and removes attached algae, and also feeds on benthic organisms and detritus, this introduced catfish may be having a significant impact on the aquatic food base and may, therefore, negatively affect native invertebrate and vertebrate species.

100

Male members of the genus Pterygoplichthys dig out river banks to create burrows in which an attracted female will lay and guard her eggs. In large numbers, this burrowing behavior by Pterygoplichthys contributes to problems with siltation. In addition, the burrows potentially destabilize the banks, leading to an increased rate of erosion (Nico and others, 2009).

Pterygoplichthys spp. may compete with native fish and are also believed to have displaced several species of native minnows in Texas, including the federally threatened Devils River minnow (Diona diaboli). These catfishes have been found to ingest eggs of Etheostoma fonticola, which is listed as vulnerable (Cohen, 2008; Cook-Hildreth, 2009).

Comments: Sailfin suckermouth catfishes (Pterygoplichthys spp.) are capable of surviving mesohaline conditions (up to 10 ppt) for extended periods of time, allowing for the use of estuarine and coastal areas for dispersal (Capps and others, 2011). Several armored catfish, identified as this species by J. Armbruster, were taken in Texas from a water body in the Galveston Bay Basin near Houston in 1998 (Nico and Martin, 2001; museum specimen); a single small specimen was preserved.

Osteoglossum bicirrhosum (Cuvier, 1829) (Arawana)

Habitat: The Arawana inhabits shallow, blackwater lagoons and littoral zones of rivers and lakes of Amazonia. It also invades the flooded forest during annual high-water periods (Saint-Paul and others, 2000).

Life History: The Arawana is a generalist predator and although it will take fish and other vertebrates, it feeds primarily on insects and other arthropods (Goulding, 1980). An Arawana may leap from the water to feed on terrestrial or arboreal prey (Goulding, 1980; Lowry and B. Loftus, U.S. Geological Survey others, 2005). This species is a mouthbrooder, with males performing the brooding role (Goulding, 1980).

Native Range: Tropical America; northern South America including the Amazon Basin and the Guianas (Robins and others, 1991).

Nonindigenous Range: A fish was photographed at Blue Hole on Big Pine Key, Florida, in 2003.

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Impacts: Unknown.

Comments: Conventionally, most Arawanas (especially in the aquarium trade) have been referred to as O. bicirrhosum; however, some of these (including USGS records) may have actually been O. ferreirai.

Polypterus delhezi Boulenger, 1899 (Barred Bichir)

Habitat: The Barred Bichir lives in streams, lakes, and flood zones (Matthes 1964 in Froese and Pauly, 2012).

Life History: Spawning occurs during the rainy season. This species is carnivorous. Polypterus delhezi is a demersal, air breathing species (Matthes 1964 in Froese and Pauly, 2012).

Native Range: Polypterus delhezi is native to the Congo River Basin in Africa (Gosse, 1984). Steven G. Johnson

Nonindigenous Range: Several small (approximately 100 mm TL) individuals were collected from a drainage canal near Coral Springs, Broward County, Florida, in 2001 (Shafland and others, 2008b).

102

Geography Native Range Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown

Vector: Aquarium release.

Impacts: Unknown.

Tetraodon nigroviridis (Marion de Procé, 1822) (Spotted Green Pufferfish)

Habitat: The species lives in fresh- or brackish-water areas. In its native habitat, the Spotted Green Pufferfish inhabits streams, rivers, and floodplains (Rainboth, 1996).

Life History: The Spotted Green Pufferfish feeds on mollusks, crustaceans, other vertebrates, and some plant matter (Rainboth, 1996).

Native Range: Tetraodon nigroviridis is native to Asia, from Sri Lanka to Indonesia and north to China (Froese and Pauly, Shannon Ellis 2012).

Nonindigenous Range: Three specimens were collected in a tributary to Big Branch Bayou, St. Tammany Parish, Lacombe, Louisiana, in 2004.

Vector: Aquaculture escape from a nearby tropical fish farm. Geography Native Range Impacts: Unknown. Non-specific State Introduction Introduced Range Site Accuracy Accurate Approximate Centroid Population Status Established Failed Unknown 103

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