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The Nonindigenous Occurrences section of the NAS species profiles has a new structure. The section is now dynamically updated from the NAS database to ensure that it contains the most current and accurate information. Occurrences are summarized in Table 1, alphabetically by state, with years of earliest and most recent observations, and the tally and names of drainages where the species was observed. The table contains hyperlinks to collections tables of specimens based on the states, years, and drainages selected. References to specimens that were not obtained through sighting reports and personal communications are found through the hyperlink in the Table 1 caption or through the individual specimens linked in the collections tables.

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Common name: Coqui

Synonyms and Other Names: Puero Rican coqui, coquí común

Taxonomy: available through www.itis.gov

Identification: A small, brown or gray-brown arboreal frog which, unlike the nonindigenous greenhouse frog (Eleutherodactylus planirostris), has eyes that are gold, golden-brown, or brown, rather than red, and has toe disks (toepads) for climbing (Conant and Collins, 1998; Joglar, 1998; Rivero, 1998).  May superficially resemble indigenous Floridian treefrogs.  Dorsal pattern varies considerably ranging from no pattern, to one or two broad cream stripes, v-shaped marks, spots, botches, or an ill-defined pale band or “M” between the shoulders (illustrated in Bartlett, 1994; Rivero, 1998; Bartlett and Bartlett, 1999; and described in Powell et al., 1998). 

Call of male coqui is a rapid, loud “ko-KEE”, hence its common name (Bartlett, 1994; Conant and Collins, 1998; Rivero, 1998; Bartlett and Bartlett, 1999; Thomas, 2006). A recording of the call of the coqui is available on a CD by Rivero (1998) and online (Thomas, 2006).

Size: 33-57 mm snout-rear length with Florida populations averaging slightly smaller, typically not exceeding 44.4 mm (Bartlett and Bartlett, 1999).

Native Range: Native to Puerto Rico in a wide variety of habitats and elevations (Joglar and Burrowes, 1996; Powell et al., 1996; Stewart, 1996; Powell, 1999; Thomas, 1999).

Alaska	Hawaii	Puerto Rico & Virgin Islands	Guam Saipan

Hydrologic Unit Codes (HUCs) Explained
Interactive maps: Point Distribution Maps

Ecology: These treefrogs are nocturnal insectivores, and groups can consume up to 114,000 invertebrates per hectare in a single evening (Stewart and Woolbright, 1996). They are habitat generalists, and do not show preference for forest floor height or plant species (Beard et al., 2003). Coqui are highly fecund and in exist in the largest densities known by any amphibian in the world, and have been recorded at up to 20,000 individuals per hectare (Stewart, 1995; Kraus and Campbell, 2002; Woolbirght et al. ,2006). Near the Hilo area of eastern Hawaii Island (Big Island), population densities of E. coqui are three times higher than those in their native Puerto Rico (Woolbright et al., 2006). Coqui may reach the ground from their arboreal perches by parachuting (Stewart, 1985).  To attract a mate, males call to females, with the greater mating advantage going to the males with the highest call rates (Townsend and Stewart, 1986a; Lopez, 1996). Calls also are used in aggressive encounters (Stewart and Rand, 1991, 1992).

Fertilization is internal; development is direct with well-developed neonate frogs hatching from eggs in vegetation without any aquatic tadpole stage (Townsend et al., 1981; Townsend and Stewart, 1985, 1986b; Townsend, 1996).  Males brood eggs in an elevated, sheltered spot such as a cavity or a bromeliad (Taigen et al., 1984; Townsend et al., 1984; Townsend, 1986).  In Hawaii, E. coqui supplement their shelter sites and nesting sites by making extensive use of subterranean passages and galleries within the porous lava substrate (Woolbright et al., 2006).

Means of Introduction: Most introductions are probably the result of horticultural and landscaping imports; many plants provide shelter for both the adult frogs and their eggs (Austin and Schwartz, 1975; Kraus et al., 1999; Kraus and Campbell, 2002), although some releases in Hawaii are intentional (Woolbright et al., 2006; F. Kraus, personal communication 2002). Everman and Klawinski (2013) examined the population genetics and dispersal patterns of Coqui populations on Hawai'i Island, finding no relationship between genetic and geographic distance, suggesting that the jump-dispersal pattern exhibited by this species is accomplished through human-mediated assistance.

Status: Status populations in the U.S. Virgin Islands remain established (Schwarz and Henderson, 1991; Conant and Collins, 1998). An erroneous Louisiana record is based on two pet coqui, both males, kept in a greenhouse for 2-3 years until killed off by a winter freeze (Dundee, 1991). Populations in Miami-Dade County, Florida, persist and are limited to areas in and around a few greenhouses (Loftus and Herndon, 1984; Bartlett, 1994; Meshaka et al., 2004).  Those coqui outside the greenhouses tend to die off during winter freezes and those once found at the Fairchild Tropical Gardens are now suspected extirpated (Wilson and Porras, 1983; Ashton and Ashton, 1988; Bartlett and Bartlett, 1999).  It is not known if the Florida populations are self-sustaining or replenished through new horticultural plantings (Bartlett and Bartlett, 1999).

Numerous populations of coqui in Hawaii are established, and considered highly invasive (Hawaii Invasive Species Council, 2018). A variety of methods are being used to monitor and eradicate these rapidly spreading frogs (Kraus et al., 1999; Kraus and Campbell, 2002; Thomas, 2006; Woolbright et al., 2006).

Impact of Introduction: Impact of E. coqui in the U.S. Virgin Islands remains unknown, but due to the similarity in fauna and habitat so relatively close to their native range and ecology, Kraus et al. (1999) expect few problems. The impact of the few Florida populations remains unknown. Hawaii has no native frogs, and Hawaiian populations of these insectivores are spreading rapidly and considered injurious wildlife by the Hawaii Invasive Species Council (2018) (Kraus et al., 1999; Woolbright et al., 2006). Coqui could potentially eat indigenous, endemic arthropods, including species of insects and spiders close to extinction (Kraus et al., 1999). This also could have a negative impact on indigenous insectivorous birds that may be forced to compete with E. coqui for food (Kraus et al., 1999; Thomas, 2006).  Nutrient flow through the native food web may be disrupted, and coqui may serve as a source of food for nonindigenous, invasive predators (Kraus, 1999; Woolbright et al., 2006). Coqui served as a food source for local birds in Hawaii, rather than a competitor for insects (Smith, 2016). While non-native mongoose (Herpestes auropunctatus) in Hawaii eat coqui and as a result have higher abundance than in areas without coqui, non-native Pacific Rats (Rattus exulans) have lower abundance in Coqui-invaded areas, apparently as a result of competition for invertebrate food (Hill et al. 2019).

In Hawaii, the loud calls of the coqui have caused residents to loss sleep, tourists have lodged complaints with hotels, and residents may have difficulty selling infested property or experience weaker property values (Kraus et al., 1999; Kraus and Campbell, 2002; Thomas, 2006). Woolbright et al. (2006), recorded sound pressure levels of calling male coqui in eastern Hawaii Island (Big Island) up to 73 dB. Further human concerns are discussed in detail by Kraus and Campbell (2002).

Remarks: The taxonomy and taxonomic placement of coqui is reviewed in Frost (2000) and Frost et al. (2006). Over the years E. coqui in their native range have been the subject of a wide variety studies dealing with ecology, behavior and reproduction (e.g., Townsend and Mogler, 1987; Woolbright, 1989; Beard et al., 2003).

There is current concern that E. coqui may become established in Guam in the same fashion as the closely related greenhouse frog, E. planirostris (Hurley, 2003); they have been reported in Guam, but their status is currently unknown.  See the species account titled “Eleutherodactylus planirostris (Cope, 1862)” on this website.

In their native Puerto Rico, populations in cloud forests are on the decline, while those populations found in other habitats remain stable (Joglar and Burrowes, 1996).

References: (click for full references)

Ashton, R. E., Jr., and P. S. Ashton. 1988. Handbook of Reptiles and Amphibians of Florida. Part Three. The Amphibians. Windward Publishing, Inc., Miami. 191 pp.

Austin, D. F., and A. Schwartz. 1975. Another exotic amphibian in Florida, Eleutherodactylus coqui. Copeia 1975(1):188.

Bartlett, R. D. 1994. Florida’s alien herps.  Reptile & Amphibian Magazine (27):56-73, 103-109.

Bartlett, R. D., and P. P. Bartlett. 1999. A Field Guide to Florida Reptiles and Amphibians. Gulf Publishing Company, Houston. 280 pp.

Beard, K. H., S. McCullough, and A. K. Eschtruth. 2003. Quantitative assessment of habitat preferences for the Puerto Rican terrestrial frog, Eleutherodactylus coqui. Journal of Herpetology 37(1):10-17.

Butterfield, B. P., W. E. Meshaka, Jr., and C. Guyer. 1997. Nonindigenous amphibians and reptiles.  Pp. 123-138. In: D. Simberloff, D. C. Schmitz, and T. C. Brown (editors). Strangers in Paradise. Impact and Management of Nonindigenous Species in Florida. Island Press, Washington, D. C. 467 pp.

Conant, R., and J. T. Collins. 1991. A Field Guide to Reptiles and Amphibians. Eastern and Central North America. Third Edition. Houghton Mifflin Company, Boston. 450 pp.

Conant, R., and J. T. Collins. 1998. A Field Guide to Reptiles & Amphibians. Eastern and Central North America. Third Edition, Expanded. Houghton Mifflin Company, Boston. 616 pp.303 pp.

Dundee, H. A. 1991. When is an introduction not an introduction? Herpetological Review 22(4):122.

Everman, E., and P. Klawinski. 2013. Human-facilitated jump dispersal of a non-native frog species on Hawai'i Island. Journal of Biogeography 40:1961-1970. http://onlinelibrary.wiley.com/doi/10.1111/jbi.12146/abstract.

Frost, D. [R.] (compiler). 2000. Anuran frogs. Pp. 6-17. In: B. I. Crother (chair), and Committee on Standard English and Scientific Names (editors). Scientific and standard English names of amphibians and reptiles of North America north of Mexico, with comments regarding confidence in our understanding. Society for the Study of Amphibians and Reptiles Herpetological Circular (29):i-iii, 1-82.

Frost, D. R., T. Grant, J. Faivovich, R. H. Bain, A. Haas, C. F. B. Haddad, R. O. De Sá, A. Channing, M. Wilkinson, S. C. Donnellan, C. J. Raxworthy, J. A. Campbell, B. L. Blotto, P. Moler, R. C. Drewes, R. A. Nussbaum, J. D. Lynch, D. M. Green, and W. C. Wheeler. 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History 297:1-370 + Fig. 50 foldout.Henderson, R. W., and R. Powell. 1999. West Indian herpetoecology. Pp. 223-268. In: B. I. Crother (editor). Caribbean Amphibians and Reptiles. Academic Press, San Diego. 495 pp.

Hawaii Invasive Species Council. 2018. Coqui frog (Eleutherodactylus coqui). https://dlnr.hawaii.gov/hisc/info/invasive-species-profiles/coqui/. Created on 02/21/2013. Accessed on 05/02/2018.

Hill, S.A., K.H. Beard, S.R. Siers, and A.B. Shiels. 2019. Invasive coqui frogs are associated with differences in mongoose and rat abundances and diets in Hawaii. Biological Invasions 21(6):2177-2190. https://link.springer.com/article/10.1007/s10530-019-01965-3.

Hurley, T. 2003. Frog find fuels snake concern. HonoluluAdvertiser.com [online] 2003(12 November). Available on URL: http://the.honoluluadvertiser.com/article/2003/Nov/12/ln/ln09a.html.

Joglar, R. L. 1998. Los Coquíes de Puerto Rico. Su Historia Natural y Conservación. Editorial de las Universidad de Puerto Rico, San Juan. 232 pp.

Joglar, R. L., and P. A. Burrowes. 1996. Declining amphibian populations in Puerto Rico. Pp. 371-380. In: R. Powell and R. W. Henderson (editors). Contributions to West Indian Herpetology. A Tribute to Albert Schwartz. Contributions to Herpetology 12. Society for the Study of Amphibians and Reptiles, Ithaca. 475 pp.

Joglar, R. L., and N. Rios-López. 1998. Geographic distribution: Eleutherodactylus coqui (Puerto Rican coqui). Dominican Republic: Districto Nacional. Herpetological Review 29(2):107.

King, F. W. 2006. Checklist of Florida Amphibians and Reptiles [online]. Florida Museum of Natural History, University of Florida, Gainesville. Available on URL: http://www.flmnh.ufl.edu/herps/FL-GUIDE/Flaherps.htm.

Kraus, F. 2002. Personal communication—Zoologist, Bishop Museum, 1525 Bernice Street, Honolulu, Hawaii.

Kraus, F., and E. W. Campbell III. 2002. Human-mediated escalation of a formerly eradicable problem: The invasion of Caribbean frogs in the Hawaiian Islands. Biological Invasions 4(3):327-332.

Kraus, F., E. W. Campbell, A. Allison, and T. Pratt. 1999. Eleutherodactylus frog introductions to Hawaii. Herpetological Review 30(1):21-25.

Loftus, W. E., and R. Herndon. 1984. Reestablishment of the coqui, Eleutherodactylus coqui Thomas, in southern Florida. Herpetological Review 15(1):23.

Lopez, P. T. 1996. Mate selection in the Puerto Rican frog, Eleutherodactylus coqui. Pp. 241-250. In: R. Powell and R. W. Henderson (editors). Contributions to West Indian Herpetology. A Tribute to Albert Schwartz. Contributions to Herpetology 12. Society for the Study of Amphibians and Reptiles, Ithaca. 475 pp.

McCann, J. A., L. N. Arkin, and J. D. Williams. 1996. Nonindigenous Aquatic and Selected Terrestrial Species in Florida. Status, Pathways, Dates of Introduction, Range Distributions, and Significant Ecological and Economic Effects. Florida Caribbean Science Center, U. S. Geological Survey, Gainesville. 301 pp.

McCoid, M. J., and C. Kleberg. 1995. Non-native reptiles and amphibians. Pp. 433-437. In: E. T. LaRoe, G. S. Farris, C. E. Puckett, P. D. Doran, and M. J. Mac (editors). Our Living Resources: A Report to the Nation on the Distribution, Abundance, and Health of U. S. Ecosystems. U.S. Department of the Interior, National Biological Service, Washington, D. C. 530 pp.

McKeown, S. 1998. Notes on a newly established frog, Eleutherodactylus coqui, in the Hawaiian Islands. Bulletin of the Chicago Herpetological Society 33(2):30-31.

Meshaka, W. E., Jr., B. P. Butterfield, and J. B. Hauge. 2004.  The Exotic Amphibians and Reptiles of Florida. Krieger Publishing Company, Malabar, Florida. 166 pp.

Moler, P. 1988. A Checklist of Florida’s Amphibians and Reptiles. Nongame Wildlife Program, Florida Game and Fresh Water Fish Commission, Tallahassee. 18 pp.

Powell, R. 1999. Legends to color plates.  Pp. xxi-xxx, plate 5. In: B. I. Crother (editor). Caribbean Amphibians and Reptiles. Academic Press, San Diego. 495 pp.

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Henderson (editors). Contributions to West Indian Herpetology. A Tribute to Albert Schwartz. Contributions to Herpetology 12. Society for the Study of Amphibians and Reptiles, Ithaca. 475 pp.

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Smith, R.L. 2016. Invasive Coqui Frogs Serve as Novel Prey for Birds in Hawaii, and Not as Competitors. Unpublished M.S. thesis. Utah State University, Logan, UT.

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Stewart, M. M. 1995. Climate driven population fluctuations in rain forest frogs. Journal of Herpetology 29:437-446.

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Stewart, M. M., and A. S. Rand. 1991. Vocalizations and the defense of retreat sites by male and female frogs, Eleutherodactylus coqui. Copeia 1991(4):1013-1024.

Stewart, M. M., and A. S. Rand. 1992. Diel variation in the use of aggressive calls by the frog Eleutherodactylus coqui. Herpetologica 48(1):49-56.

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Taigen, T. L., F. H. Pough, and M. M. Stewart. 1984. Water balance of terrestrial anuran (Eleutherodactylus coqui) eggs: Importance of parental care. Ecology 65(1):248-255.

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Townsend, D. S. 1986. The cost of male parental care and its evolution in a neotropical frog. Behavioral Ecology and Sociobiology19:187-195.

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Townsend, D. S., and W. H. Moger. 1987. Plasma androgen levels during male parental care in a tropical frog. Hormones and Behavior 21:93-99.

Townsend, D. S., and M. M. Stewart. 1985. Direct development in Eleutherodactylus coqui (Anura: Leptodactylidae): A staging table. Copeia 1985(2):423-436.

Townsend, D. S., and M. M. Stewart. 1986a. Courtship and mating behavior of a Puerto Rican frog, Eleutherodactylus coqui. Herpetologica 42(2):165-170.

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Townsend, D. S., M. M. Stewart, F. H. Pough, and P. F. Brussard. 1981. Internal fertilization in an oviparous frog. Science 212:469-471.

Vivarium Staff. 1998. Hawaii’s war on herps. Vivarium 6(6):6, 8, 17, 61.

Walls, J. G. 1998. Say goodbye to Jackson’s chameleons? Reptile Hobbyist 4(3):92-93.

Wilson, L. D., and L. Porras. 1983. The ecological impact of man on the South Florida herpetofauna. University of Kansas Museum of Natural History Special Publication (9):i-vi, 1-89.

Woolbright, L. L. 1989. Sexual dimorphism in Eleutherodactylus coqui: Selection pressures and growth rates. Herpetologica 45(1):68-74.

Woolbright, L. L., A. H. Hara, C. M. Jacobsen, W. L. Mautz, and F. L. Benevides, Jr. 2006. Population densities of the coquí, Eleutherodactylus coqui (Anura: Leptodactylidae) in newly invaded Hawaii and in native Puerto Rico. Journal of Herpetology 40(1):122-126.

Other Resources:
Alien Frogs in Hawaii (HEAR Project)

Eradicators Concede Big Island To Frogs

$7 million sought to fight coqui frogs

Eluetherodactylus coqui (Puerto Rican coqui) (Gulf of Mexico Program)

Eluetherodactylus coqui (Global Invasive Species Database)

US Fish and Wildlife Service Ecological Risk Screening Summary for Eleutherodactylus coqui

Author: Louis A. Somma, and Matthew Neilson

This information is preliminary or provisional and is subject to revision. It is being provided to meet the need for timely best science. The information has not received final approval by the U.S. Geological Survey (USGS) and is provided on the condition that neither the USGS nor the U.S. Government shall be held liable for any damages resulting from the authorized or unauthorized use of the information.