Rana catesbeiana (American bullfrog)

Pictures

Picture	Title	Caption	Copyright
Title Adult Caption Adult American bullfrog, (Rana catesbeiana). Chodikee Lake, Ulster County, New York, USA Copyright ©Russ Ottens/University of Georgia, Bugwood.org/Creative Commons Attribution 3.0 License. CC BY 3.0 US	Adult	Adult American bullfrog, (Rana catesbeiana). Chodikee Lake, Ulster County, New York, USA	©Russ Ottens/University of Georgia, Bugwood.org/Creative Commons Attribution 3.0 License. CC BY 3.0 US
Title Swabbing Caption Chytridiomycosis; swabbing a North American bullfrog, farmed for the food trade in China, to test for Batrachochytrium dendrobatidis infection. Copyright ©Lisa M. Schloegel	Swabbing	Chytridiomycosis; swabbing a North American bullfrog, farmed for the food trade in China, to test for Batrachochytrium dendrobatidis infection.	©Lisa M. Schloegel
Title Swabbing Caption Chytridiomycosis; swabbing the hindfoot webbing of a North American bullfrog. to test for infection by the fungus Batrachochytrium dendrobatidis. Copyright ©Lisa M. Schloegel	Swabbing	Chytridiomycosis; swabbing the hindfoot webbing of a North American bullfrog. to test for infection by the fungus Batrachochytrium dendrobatidis.	©Lisa M. Schloegel
Title North American bullfrogs in culture Caption Chytridiomycosis; North American bullfrogs in Taiwan, farmed for the international food trade. Copyright ©Lisa M. Schloegel	North American bullfrogs in culture	Chytridiomycosis; North American bullfrogs in Taiwan, farmed for the international food trade.	©Lisa M. Schloegel
Title North American bullfrogs in culture Caption Chytridiomycosis; North American bullfrogs in Taiwan, farmed for the international food trade. Copyright ©Lisa M. Schloegel	North American bullfrogs in culture	Chytridiomycosis; North American bullfrogs in Taiwan, farmed for the international food trade.	©Lisa M. Schloegel
Title North American bullfrogs in culture Caption Chytridiomycosis; close view of north American bullfrogs in Taiwan, farmed for the international food trade. Copyright ©Lisa M. Schloegel	North American bullfrogs in culture	Chytridiomycosis; close view of north American bullfrogs in Taiwan, farmed for the international food trade.	©Lisa M. Schloegel

Identity

Preferred Scientific Name

• Rana catesbeiana Shaw, 1802

Preferred Common Name

• American bullfrog

Other Scientific Names

• Lithobates (Aquarana) catesbeianus Dubois, 2006
• Lithobates catesbeianus Dubois, 2006
• Rana (Aquarana) catesbeiana Dubois, 1992
• Rana (Aquarana) catesbeiana Hillis, 2007
• Rana (Lithobates) catesbeiana Shaw, 1802
• Rana (Novirana) catesbeiana Hillis and Wilcox, 2005

International Common Names

• English: bullfrog; common bullfrog; North American bullfrog
• Spanish: rana mugidora; rana toro; rana toro Americana
• French: grenouille d'Amérique; grenouille taureau; grenouille-taureau americaine; ouaouaron

Local Common Names

• Brazil: rã-touro
• Germany: ochsenfrosch
• Italy: rana toro
• Netherlands: grote kikker
• Sweden: oxgroda

Summary of Invasiveness

The American bullfrog has been transported around the world primarily with the intention of cheaply cultivating its large and meaty hind legs for human consumption. It is also of commercial interest to biological supply houses, fish bait suppliers, the pet trade, and pond landscapers. In North America in the early 20^th Century it was released outside of its natural range by wildlife agencies keen on introducing a new game species (Lannoo, 1996; Tangley, 2003). Once liberated it is extraordinarily prolific, prone to migration, and highly adaptable to new environments. It grows into a relatively large, voracious, and indiscriminate predator that can come to dominate the margins of lakes and ponds. It competes with native anurans. Larvae can have a significant impact upon benthic algae, and thus perturb aquatic community structure. Adults may be responsible for significant levels of predation on native anurans and other aquatic herpetofauna, such as snakes and turtles. The species also contributes to the spread of pathogens such as the chytrid fungus, Batrachochytrium dendrobatidis (Garner et al., 2006). R. catesbeiana has been identified by the World Conservation Union (IUCN) as one of the world’s 100 worst invasive alien species (ISSG, 2005).

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Metazoa
•         Phylum: Chordata
•             Subphylum: Vertebrata
•                 Class: Amphibia
•                     Order: Anura
•                         Family: Ranidae
•                             Genus: Rana
•                                 Species: Rana catesbeiana

Notes on Taxonomy and Nomenclature

The American bullfrog is a member of the nearly cosmopolitan family Ranidae. For most of the past 200 years this species has been unambiguously assigned to the genus Rana and the species R. catesbeiana, Shaw, 1802. Over the past 30 years, however, advances in biochemical systematics have led to both increased knowledge and increased contention concerning how new and ever-more refined data sets should be interpreted and taxonomically expressed. Thus, there are today numerous taxonomic synonyms for R. catesbeiana, e.g. Lithobates catesbeianus,Aquarana catesbeiana, and Novirana catesbeiana, that have received varying degrees of acceptance amongst herpetological systematists and the broader zoological community (Frost 2009).

Description

R. catesbeiana is not the largest frog species in the world but it is one of the top ten (and the largest true frog in North America) with a maximum body length slightly in excess of 200 mm (typical length 90-152 mm) and body weight up to 0.5 kg. Like most frogs, it undergoes a drastic metamorphosis during its life cycle, passing from a young aquatic life phase with branchial respiration, predominantly plankton feeding, iliophagous or herbivorous, to reach adult life as an animal with pulmonary and skin respiration and a carnivorous feeding habit (Teixeira et al., 2001). Bullfrog tadpoles are also very large by frog standards (80-150 mm) and can take from 12 to 48 months to reach metamorphosis. A bullfrog tadpole’s body can be as large as a golf ball with a relatively long, high-finned and muscular tail. The colouration of the tadpole stage is brown to light olive with small black spots scattered across the head and upper body. At metamorphosis the tadpoles resorb their gills and finned tails while transforming into juvenile miniatures of adult bullfrogs but without secondary sexual characters. The colour of adults varies from olive, green or brownish on the dorsum with vague spots or blotches; the head is lighter green, and the legs blotched or banded; the eardrums are conspicuous. The hind feet are fully webbed. The skin is mostly smooth. There are no dorsolateral folds; a short fold extends from the eye over and past the eardrum to the forearm.

Distribution

R. catesbeiana is native to eastern North America, ranging naturally from Nova Scotia, southern Quebec and Ontario in Canada, down through the eastern United States and Mississippi drainage, and southward along the east coast of Mexico. It has been introduced to Hawaii, parts of the western USA and Canada, Mexico and the Caribbean, South America, Europe and Asia (ISSG, 2005). In the central United States it is difficult to say for certain where natural populations end and alien populations begin, but there is no disputing the fact that all occurrences west of the Rocky Mountains are the result of translocation and release during the late 19^th century and throughout the 20^th. (It is known that bullfrogs were introduced to areas of California and Colorado in the early 1900s -- University of Michigan Museum of Zoology, 2005).

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

History of Introduction and Spread

In the late 19^th and early 20^th centuries bullfrogs were translocated from the eastern United States to many western states (Moyle, 1973; Bury and Whelan, 1984) and Hawaii (Pitt et al., 2005), western Canada (Orchard, 1999), the Caribbean (Kairo et al., 2003; Kraus, 2009), western Europe (Ficetola et al., 2007 -- see electronic appendix to this paper for a table with details of introductions to several European countries, mostly in the late 20th century; Lanza, 1962; Veenvliet and Veenvliet, 2003, 2004; Nehring and Klingenstein, 2008), South America (Hanselmann et al., 2004; Giovanelli et al., 2008; Laufer et al., 2008; Kraus, 2009) east Asia (Fei et al., 1999; Hirai, 2004; Wu et al., 2005; Wang et al., 2008) and southeast Asia (Hardouin, 1997). There have also been introductions to western and central Mexico, from the eastern USA and north-eastern Mexico.

Introductions

Risk of Introduction

People continue to routinely transport bullfrogs from one place to another because they appear to be an easily cultivated and profitably marketable commodity for human consumption. Bullfrogs almost inevitably escape because they are, if not impossible, then certainly impractical to confine (see Liu and Lee (2009) for information on the effects of enclosure type and other factors on the risk of escape). However even in the face of wide recognition that it is not a good thing to do, some people persist in moving this ecologically damaging species from place to place for a complex of reasons that range from genuine ignorance to wilful mischief.      

Habitat

Bullfrogs will colonize a wide variety of lakes, ponds, reservoir, irrigation ditches and marshes, but there are vital characteristics for supporting a population. Permanent water is necessary because bullfrog tadpoles generally take at least 12 months and as much as 48 months to reach metamorphosis; seasonal or intermittent pools, though useful as way stations for migrating bullfrogs, will probably not permit successful reproduction. Water temperature is also important because bullfrogs reproduce only in very warm water when summer temperatures exceed 25 degrees Celsius. For this reason, lakes and ponds that are not too deep < 6 m) and have good sun-exposure around the margins provide most favourable habitat for bullfrog reproduction. Riparian thickets and an abundance of aquatic and emergent vegetation are other factors that, while not vital, are often associated with thriving bullfrog populations because they provide cover and likely a more diverse prey base. However, golf course ponds with edges denuded of most riparian plants can also support healthy bullfrog populations. Acidification of waterways has been associated with bullfrog population declines in southern Ontario (Berrill et al., 1992), but the tolerance limits of bullfrogs to water chemistry extremes and various synthetic chemicals are still being explored.

Habitat List

Biology and Ecology

Genetics

Recent studies have looked at genetic markers to try to identify the geographical origin, number of founders, and number of introductions of alien, invasive bullfrog populations (Ficetola et al, 2008a; Austin et al., 2004). While this line of research has demonstrated success at achieving its scientific goal, it has not yet been applied to bullfrog management and eradication. These studies do highlight the fact that American bullfrogs are physiologically adaptable and behaviourally flexible when moved between environments that are markedly dissimilar, and that they can invade huge areas starting from only a small number of founder individuals.

Natural Food Sources

Climate

Latitude/Altitude Ranges

Water Tolerances

Natural enemies

Notes on Natural Enemies

Bullfrog eggs are commonly attacked by fungi (Ruthig, 2009) and perhaps leeches (Licht 1969), while the tadpoles are preyed upon by aquatic invertebrates such as dragonfly larvae and predatory water beetles. Although fish sometimes predate bullfrog eggs, tadpoles and juveniles (Bury and Whelan, 1984; Casper and Hendricks, 2005), tadpoles avoid predation by being distasteful to many of the more widespread predatory fish, or use chemical-cued avoidance behaviour in their presence (Pearl et al., 2003). Chemical-cued avoidance behaviour is also used by bullfrog tadpoles in the presence of predatory dragonfly nymphs (Peacor, 2006). Larger aquatic-foraging garter snakes (Thamnophis spp.) might be effective predators of bullfrog tadpoles and juveniles, but garter snakes are commonly eaten by adult bullfrogs. Avian predators include herons and egrets (Ardeidae; Casper and Hendricks, 2005); various reptiles and mammals also eat bullfrogs.

Means of Movement and Dispersal

Bullfrog tadpoles will spread throughout the water bodies in which they have hatched, though they tend to seek out the warmest water. From here they can enter the inflow and outflow waterways that feed and drain lakes and ponds and therefore have a limited ability to migrate. However, because bullfrog larvae (tadpoles) are gill-breathing and entirely aquatic this life stage is unable to move out of the water on its own. 

Pathway Causes

Pathway Vectors

Impact Summary

Economic Impact

Environmental Impact

Bullfrogs are prolific and aggressive competitors for space and voracious predators of a very wide variety of organisms, so displacement of native species is the primary problem that they create (Bury and Whelan, 1984; Lannoo, 2005; Santos-Barrera et al., 2009). They have a much higher critical thermal maximum than most other frogs, meaning that they are able to thrive in higher water temperature, and have a longer breeding season and a higher rate of pre-metamorphic survivorship, which also allows them to be more successful than other frogs. (They also do well with changes in the environment that have occurred due to human modification).

Threatened Species

Social Impact

There have been cases of severe allergic reaction in some people who ingest the meat of bullfrogs (Hilger et al.,2002). Also, bullfrog tadpoles and metamorphs have been shown to be suitable hosts for the pathogenic bacterium Escherichia coli (Gray et al., 2007). In Japan it was found that 92% of the bullfrogs sampled were highly infected with Blastocystis, a single-celled parasite that infects the gastrointestinal tract of hosts including humans (Yoshikawa et al., 2004). The skinning of bullfrogs has been implicated in rare cases of nematode infection of humans (Quirks and Quarks, CBC Radio, Canadian Broadcasting Corporation). Studies are currently under way looking into whether bullfrogs could play a role in transmission of West Nile virus (WNV), because the virus has been isolated in many amphibian-feeding species of mosquitoes (Klenk and Komar, 2003; Danner & Phillips, 2008). An investigation of a cholera outbreak in Hunan, China, in 2006 concluded that aquatic products such as snapping turtles and bullfrogs constituted the major causes of cholera (Deng et al., 2008).

Risk and Impact Factors

• Invasive in its native range
• Proved invasive outside its native range
• Has a broad native range
• Abundant in its native range
• Highly adaptable to different environments
• Is a habitat generalist
• Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
• Pioneering in disturbed areas
• Capable of securing and ingesting a wide range of food
• Highly mobile locally
• Benefits from human association (i.e. it is a human commensal)
• Long lived
• Fast growing
• Has high reproductive potential

• Altered trophic level
• Conflict
• Damaged ecosystem services
• Ecosystem change/ habitat alteration
• Negatively impacts aquaculture/fisheries
• Reduced native biodiversity
• Threat to/ loss of endangered species
• Threat to/ loss of native species

• Pest and disease transmission
• Predation
• Rapid growth

• Highly likely to be transported internationally deliberately
• Highly likely to be transported internationally illegally
• Difficult/costly to control

Uses

In many parts of the world where American bullfrogs are non-native, the adults are known to be hunted for food, e.g. in China. It is not known whether this utilization of bullfrogs provides an important dietary supplement or an important source of income, or is simply a recreational activity.

Uses List

Animal feed, fodder, forage

• Bait/attractant

General

• Botanical garden/zoo
• Capital accumulation
• Laboratory use
• Pet/aquarium trade
• Research model
• Sport (hunting, shooting, fishing, racing)

Human food and beverage

• Fresh meat
• Meat/fat/offal/blood/bone (whole, cut, fresh, frozen, canned, cured, processed or smoked)

Similarities to Other Species/Conditions

The average person might find it difficult to distinguish between many species in the widely distributed and speciose family Ranidae (853 described species) because so many of them are green, brown and/or olive-coloured with black irregular patterns. In any case, colour and pattern are not the most reliable of bullfrog characteristics. Size narrows the possibilities where large adult specimens are concerned – a very big specimen is likely to be a bullfrog -- but juvenile bullfrogs can easily be mistaken for other species. The low, sonorous timbre of a male bullfrog’s advertisement call is distinctive compared to that of other frogs. It sounds more in the vocal range of a cow than a frog and amounts to multiple repetitions of “Rrrungh” that vary in volume and intensity depending upon the size of the frog and the temperature of the water in which it is sitting. Juvenile bullfrogs have a distinctive escape behaviour. They produce a characteristic “MEEP” alarm call as they propel themselves into the water when frightened. This action commonly involves skipping over the water surface by rapid multiple kicks of the hind legs before submerging and swimming away.  

Prevention and Control

Prevention

Gaps in Knowledge/Research Needs

There is a nearly universal agreement that American bullfrogs are one of the worst alien invasive species internationally. They are ecologically destructive and therefore eradication is acknowledged as the most desirable goal. However, there is also a collective sense of defeatism permeating all recent published discussions concerning bullfrog management (Adams and Pearl, 2007; Kraus, 2009). Adams and Pearl (2007) conclude that the “lack of obvious economic impacts” and a lack of reasonably feasible control methods inevitably translate into a lack of political will to supply the resources for large-scale management. Kraus (2009) lists some apparently successful eradication case studies that have been carried out in Europe, but then concludes that these may be exceptions to the general pattern of eradication failure. However, Kraus also observes that large-scale ecological problems are beyond the capabilities and outside the skill sets of the average volunteer organization or institutional research lab, though these are most often the people who are expected to take the initiative. In fact, eradication programs are more akin to military operations. The strategies, tactics and techniques for an effective eradication are to out-flank, out-pace, and ultimately to overwhelm your foe. Traditionally, biologists are trained to study a problem, collect data, and publish results but eradication is something quite different in the degree of dedication and improvisation required. It also needs sufficient long-term commitments from funding agencies as well as adequate support staff.      

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Links to Websites

Organizations

World: Invasive Species Specialist Group (ISSG), Web based, http://www.issg.org/contact.htm

World: IUCN/SSC Global Amphibian Assessment, IUCN/SSC, International, http://www.issg.org

Italy: FAO (Food and Agriculture Organization of the United Nations), Viale delle Terme di Caracalla, 00100 Rome, http://www.fao.org/

Mexico: CONABIO - National Commission for Knowledge and Use of Biodiversity, Liga Periférico - Insurgentes Sur, Núm. 4903, Col. Parques del Pedregal, Delegación Tlalpan, 14010, http://www.conabio.gob.mx

USA: US Geological Survey - USGS, USGS National Center 12201 Sunrise Valley Drive, Reston, VA 20192, http://nas.er.usgs.gov

Contributors

08/12/2009 Original text (Invasive Species Compendium) by:

Stan Orchard, BullfrogControl.com Inc., 69A Burnside Road West, Victoria, British Columbia, V9A 1B6, Canada

04/05/2005 Original text (Aquaculture Compendium) by:

Marta Stéfani, Departamento de Zootecnia da FCAV/Unesp Jaboticabal, Via de Ac. Prof. Paulo Donato Castellane, s/n Jaboticabal - SP - 14884-900, Brazil

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