Eleutherodactylus planirostris
(greenhouse frog)

Pictures

Picture	Title	Caption	Copyright
Title Adult Caption Eleutherodactylus planirostris (greenhouse frog); adult. Mottled colour phase. Big Island, Hawaii, USA. May 2009. Copyright ©Christina Olson-2009	Adult	Eleutherodactylus planirostris (greenhouse frog); adult. Mottled colour phase. Big Island, Hawaii, USA. May 2009.	©Christina Olson-2009
Title Adult Caption Eleutherodactylus planirostris (greenhouse frog); close-up of adult head. Mottled color phase. Big Island, Hawaii, USA. May 2009. Copyright ©Christina Olson-2009	Adult	Eleutherodactylus planirostris (greenhouse frog); close-up of adult head. Mottled color phase. Big Island, Hawaii, USA. May 2009.	©Christina Olson-2009
Title Adults Caption Eleutherodactylus planirostris (greenhouse frog); two adults. Mottled colour phase. Big Island, Hawaii, USA. July 2009. Copyright ©Christina Olson-2009	Adults	Eleutherodactylus planirostris (greenhouse frog); two adults. Mottled colour phase. Big Island, Hawaii, USA. July 2009.	©Christina Olson-2009
Title Juvenile Caption Eleutherodactylus planirostris (greenhouse frog); juvenile. Striped colour phase. Mindanao Island, Philippines. April 2016. Copyright ©Christina Olson-2016	Juvenile	Eleutherodactylus planirostris (greenhouse frog); juvenile. Striped colour phase. Mindanao Island, Philippines. April 2016.	©Christina Olson-2016
Title Juvenile Caption Eleutherodactylus planirostris (greenhouse frog); juvenile. Striped colour phase. (Note US dime for scale, approx. 18mm dia.) Sarasota, Florida, USA. July 2010. Copyright ©Christina Olson-2010	Juvenile	Eleutherodactylus planirostris (greenhouse frog); juvenile. Striped colour phase. (Note US dime for scale, approx. 18mm dia.) Sarasota, Florida, USA. July 2010.	©Christina Olson-2010

Identity

Preferred Scientific Name

• Eleutherodactylus planirostris Cope

Preferred Common Name

• greenhouse frog

Other Scientific Names

• Eleutherodactylus planirostris Hedges
• Eleutherodactylus planirostris planirostris Schwartz
• Eleutherodactylus ricordii planirostris Shreve
• Euhyas planirostris Frost
• Hylodes planirostris Cope
• Lithodytes ricordii Cope

International Common Names

• English: Cuban flathead frog
• Spanish: rana-ladrona de invernadero

Summary of Invasiveness

E. planirostris is a frog native to Cuba, the Bahamas and the Cayman Islands. It has been introduced to countries worldwide and there are established populations in the Caribbean, North and Central America, Oceania and Asia. This species is commonly transported unintentionally as a hitchhiker in nursery plants and materials and can rapidly colonise new areas. E. planirostris thrives in human-altered landscapes and is well-adapted to a variety of disturbed and natural habitats. It may threaten native invertebrate communities and compete with native insectivore species. Due to its small size, inconspicuous call and habitat, initial infestations of this species are often overlooked until populations are already too large and widespread to control. E. planirostris is listed as invasive in Florida, Louisiana and Hawaii, USA.

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Metazoa
•         Phylum: Chordata
•             Subphylum: Vertebrata
•                 Class: Amphibia
•                     Order: Anura
•                         Family: Leptodactylidae
•                             Genus: Eleutherodactylus
•                                 Species: Eleutherodactylus planirostris

Notes on Taxonomy and Nomenclature

The species name E. planirostris, has undergone several revisions. Originally misidentified as E. ricordii, it was later considered a subspecies of E. ricordii (Goin, 1955) and then classified as its own species (Schwartz, 1965). Prior to 1965, records from Florida and Jamaica identify E. planirostris as E. ricordii (Olson et al., 2012b). E. goini, E. casparii and E. rogersi were once considered subspecies of E. planirostris (Schwartz, 1974; Díaz and Cádiz, 2008). More recently, it has been suggested it should be classified as Euhyas planirostris, or in the subgenus Euhyas (Hedges et al., 2008).

The Latin meaning of the scientific name refers to the lack of webbing between toes “Eleutherodactylus” ("free toes") and flat (“planum”) snout (“rostrum”). The common name comes from its association with plant nurseries, gardens and greenhouses (Schwartz and Henderson, 1991). Wright and Wright (1949) also referred to it as the as Ricord’s frog, cricket toad, Bahaman tree frog and pink-snouted frog, but this account is from when it was thought to be E. ricordii and no other sources have used these common names. Its common name in Spanish is rana-ladrona de invernadero (Díaz and Cádiz, 2008).

Description

E. planirostris is a small, terrestrial frog with a number of distinguishing characteristics; its head is as broad as body, its snout truncate and extending slightly beyond the lower jaw, it has black eye colour with a brown iris with a narrow red rim around the pupil, long, slender toes that lack webbing with small, terminal disks and white or coral red tympanum, approximately half the size of the eye (Wright and Wright, 1949). The venter is white to light grey and dorsum is tan pink to dark reddish brown (Ashton and Ashton, 1988; Bartlett and Bartlett, 2006). Two colour phases are generally described: a mottled tan and brown phase and a mottled tan and brown with two yellow dorsolateral stripes extending from the eye along the length of the body (Lynn and Grant, 1940). A third colour phase of mottled with two medium beige or orange dorsolateral dots has also been reported in Cuba (Díaz and Cádiz, 2008). There is one report of albinism (Petrovic, 1973). The male advertisement call is a series of short, irregular soft chirps (Schwartz, 1974) and is often mistaken for a cricket or bird (Wright and Wright, 1949).

In Cuba, its native range, females have a maximum snout-vent-length (SVL) of 27 mm and males 21 mm (Schwartz, 1974). In its introduced range, mean SVL is variable; mean SVL for E. planirostris is 18 mm in Jamaica (Stewart and Martin, 1980), maximum female SVL is 26.5 mm and maximum male SVL is 17.5 mm in Florida (Meshaka et al., 2004), adult female SVL ranges from 17 - 27mm and adult male ranges from 14 - 21 mm in Hawaii (Olson and Beard, 2012).

Distribution

E. planirostris is native to Cuba, the Cayman Islands and northern Bahamas (Barbour and Ramsden, 1919; Schwartz, 1974; Schwartz and Henderson, 1991; Estrada and Ruibal, 1999; Hedges, 1999; Echternacht et al., 2011; Powell and Henderson, 2012). Several early distribution records considered only the Cuban population as native (Goin, 1947; Schwartz and Henderson, 1991). Schwartz et al. (1975) stated that the Bahaman and Cayman populations originated in Cuba, but does not indicate if he believed they were introduced via natural or human-mediated dispersal. However, molecular studies and fossil records indicate that the Bahamas and Grand Cayman individuals arrived naturally and prior to human colonisation (Seidel and Franz, 1994; Heinicke et al., 2011). In Cuba, E. planirostris is one of the most widely distributed amphibians and is found throughout the island of Cuba, Isla de la Juventad and the smaller islands off the northern and southern coasts of Cuba (Powell and Henderson, 2012; González et al., 2014). In the Bahamas, their native range is limited to the northern islands of Great Bahama Bank and Little Bahama Bank and in the Cayman Islands, Grand Cayman and Cayman Brac islands (Powell and Henderson, 2012).

In the Caribbean, E. planirostris has been introduced into Great Inagua Bank in the south of the Bahamas (Schwartz, 1974, Schwartz and Henderson, 1991, Powell and Henderson, 2012). Schwartz (1974) speculated that populations found on Caicos (North Caicos) are native, but E. planirostris are now considered non-native in all of the Turks and Caicos (Lever, 2003; Reynolds and Niemiller, 2010; Reynolds, 2011; Reynolds, 2012; Powell and Henderson, 2012). Established populations are recorded on the islands of Providenciales, Grand Turk, North Caicos, Middle Caicos and Big Ambergris Key. E. planirostris is widespread throughout Jamaica, more commonly in open and disturbed habitats (Crombie, 1999; Hedges, 1999; Powell et al., 2011; Wilson, 2011). There are records for E. planirostris in Grenada (Kaiser, 1992; Hedges, 1999; Kraus et al., 1999) but some think that this is a mistake (Powell et al., 2011; Powell and Henderson, 2012). In Guadeloupe, there is a record of Eleutherodactylus cf planirostris but additional records are lacking and it is possible that it may have been a misidentification (Breuil, 2002; Lorvelec et al., 2007; Powell and Henderson, 2012). This species has also been recorded in Honduras, in San Pedro Sula in 2007, Isla de la Guanaja in 2012, La Ceiba in 2012 and Tegucigalpa in 2014 (McCranie et al., 2008; McCranie and Orellana, 2014; Solís et al., 2014a; Solís et al., 2014b) and in the neighbourhood of Cerro Ancón in Panama City, Panama in 2007. They have also been found in the neighbourhoods of Costa del Este, Dorado Lakes and Balboa, Altos del Chase and Fuentes del Fresno, Panama (Crawford et al., 2011). An abundant greenhouse population was recorded in the mangroves of Cayo Mayor, Cayos Miskitos in Nicaragua in 1992 (Villa, 2015). There is one record for an individual E. planirostris in a residential area of Paramaribo, Suriname (Ouboter and Jairam, 2012) but there is no further information. Records for E. planirostris in Mexico include the port of Veracruz and, more recently, from Yucatán peninsula, in Playa del Carmen, 2010 and Cancun, 2015 (Schwartz, 1974; Cedeño-Vázquez et al., 2014; García-Balderas et al., 2016).

On Guam, an established population was found at St. John’s School, Tumon in 2003 and later populations were found in Tamuning, Mangilao and Manengon and at the US Naval Computer and Telecommunications Station (Christy et al., 2007b, Mathies et al., 2012). This species is now found throughout the island (Olson et al., 2012b). In Asia, calling males were first detected in 2013 in a landscaped residential area of Davao City on the island of Mindanao, Philippines (Olson et al., 2014). Established populations have also been found on the islands of Cebu, Luzon and Negros (Sy et al., 2015a; Sy et al., 2015b; Sy and Salgo, 2015). In Hong Kong, sightings of E. planirostris have been reported since 2011 at the Lung Fu Shan Country Park and Campus University of Hong (HKBWS, 2013; iNaturalist, 2016; Po et al., 2016). A record from Singapore reports a single individual frog in the residential area of Sembawang in 2015 (Groenworld and Law, 2016). A photograph was taken, but no specimens were collected and there have been no additional individuals observed at the site.

A biodiversity study in southeastern Nigeria reported that several E. planirostris were found at a site in 2008 (Ukpong et al., 2012), but there is no indication whether they established, or if verification photographs or specimens were collected.

In the USA, the first record for E. planirostris is from 1863 in an unspecified location of southern Florida (Cope, 1863; Cope, 1875) and in 1889 in Key West, Monroe County (Cope, 1889). Populations were later recorded in Brevard and Miami-Dade (Dade) counties in southern Florida (Barbour, 1910). Krysko et al. (2011) determined that there are vouchered specimens from 47 Florida counties and unvouchered reports for two counties; Gadsden and Hernando. The distribution of E. planirostris is considered continuous in southern Florida and disjointed in the panhandle counties of Bay, Escambia, Franklin, Okaloosa, Leon, Gadsden and Wakulla, with most records from coastal areas (Krysko et al., 2011; Meshaka, 2011; US Geological Survey, 2016). There are also new records for De Soto County in southwest Florida, Gilchrist County in northern Florida and Gulf County in the panhandle (Thawley et al., 2012; Greene, 2014; Godwin, 2014). An individual was found alive at the Detroit Zoo in a bag of mulch shipped from Gilchrist County, Florida but it is not established in the area (Zippel, 2005). In Louisiana, E. planirostris was first documented in 1975 at the Audubon Park Zoo in New Orleans, Orleans Parish, over 500 km from the nearest population at the time (Plotkin and Atkinson, 1979). Its distribution now covers 10 parishes in residential and natural areas (Dundee et al., 1989; Dundee 1994; Williams and Wygoda, 1997; Boundy, 1998; Boundy, 2004; Elbers 2007; Liner 2007; Meshaka et al., 2009). In Alabama, the first record is for Fairhope, Baldwin County in 1982 (Carey, 1982), but there are no other records for the state until Alix et al. (2014) reported an expansion into natural areas of Baldwin County. In 2013, a population was recorded in Mobile County (McConnell et al., 2015). In Georgia, E. planirostris was found in flower pots in 1998 in Savannah, Chatham County (Winn et al., 1999) and by 2008 their distribution included areas of Camden, Chatham, Glynn, Lowndes and Thomas Counties (Jensen et al., 2008). E. planirostris is common on St. Simon Island and Brunswick, Glynn County. Records in Mississippi are for Gulfport, Harrison County from 2003, an undated record for Starkville and a record for Oktibbeha County and Ocean Springs, Jackson County from 2014 (Dinsmore, 2004; Lamb and Chatfield, 2014). In South Carolina, E. planirostris was found in two different areas of Mount Pleasant, Charleston County in 2016 (Dillman and Gibbons, 2016). This is the most northern distribution record for an established E. planirostris population. There are records of E. planirostris in Texas, Missouri, Oklahoma and Michigan (Zippel, 2005; Tipton et al., 2012; US Geological Survey, 2016). Both the Missouri and Oklahoma populations are restricted to indoors (US Geological Survey, 2016). E. planirostris was first introduced to the Pacific Basin in or before 1994 to Hawaii (Kraus and Campbell, 2002). First records were from three established populations on Hawaii Island in nurseries or nursery adjacent areas and one in a residential area of Maui (Kraus et al., 1999). E. planirostris is now found on five Hawaiian Islands: Hawaii, Kauai, Lanai, Maui, Oahu (Olson et al., 2012a; Ferreira et al., 2015).

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

E. planirostris originated on the island of Cuba and spread to adjacent islands, the Bahamas and the Caymans, via oceanic crossing, prior to human settlement (Heinicke et al., 2011). Colonisation of neighbouring Caribbean islands, including Great Inagua Bank in southern Bahamas and the Turks and Caicos, is thought to be more recent and via unintentional transport, either through the nursery or shipping trade (Schwartz and Henderson, 1991; Kraus 2009, Powell et al., 2011; Reynolds, 2011). A recent introduction to Big Ambergris Cay, Turks and Caicos, possibly in or before 2011, is thought to have arrived with construction materials from Providenciales (Reynolds, 2012).

E. planirostris was introduced to Jamaica in major port cities prior to the 1930s and thought to have been transported via the shipping industry (Stewart, 1977). This species was first recorded in Montego Bay in 1937, Kingston in 1940 and by 1943, was considered to be widespread (Lynn, 1937; Lynn, 1940; Lynn and Dent, 1943). The shipping industry may have also introduced E. planirostris to Veracruz, Mexico, but more recent introductions to the Yucatán peninsula have occurred into residential areas (Cedeño-Vazquez et al., 2014) and probably transported via the nursery trade. Recent introductions to Panama and Honduras were to residential areas and believed to have occured via the nursery trade (Crawford et al. 2011, McCranie and Orellana, 2014). Molecular analysis of specimens from Panama indicates that the E. planirostris populations originated from Florida (Crawford et al., 2011).

Analysis of the genetic diversity of the populations in Florida suggests that E. planirostris may have been introduced to the Florida Keys prior to human habitation, possibly via oceanic crossing (Heinicke et al., 2011). It has long been speculated that initial colonisation to the Florida Keys may have been natural (Wilson and Porras, 1983; Lazell, 1989) but later introductions to peninsular Florida occurred unintentionally through human assisted transport, most likely in shipments of nursery plants and materials (Goin, 1944; Goin, 1947; Heinicke et al., 2011). Records indicate a northward progression of E. planirostris up peninsular Florida and from large urban areas such as Gainesville in central Florida, Jacksonville in northeast Florida and Tallahassee in northwest Florida (van Hyning, 1933; Goin, 1944; Goin, 1947; Reichard and Stevenson, 1964). Introductions of E. planirostris into Louisiana, Georgia, Mississippi, Missouri, Oklahoma and Michigan also likely originated from nursery plants and materials shipped out of Florida (Dundee et al., 1989; Platt and Fontenot, 1995; Winn et al., 1999; Dinsmore, 2004; Zippel et al., 2005; Meshaka, 2009; US Geological Survey, 2016).

E. planirostris was first found in or adjacent to nurseries in Hawaii (Kraus et al., 1999; Kraus and Campbell, 2002) and later found in nurseries in Guam with plants shipped from Hawaii (Christy et al., 2007a). It is probable that the Hawaiian population originated from Florida (Heinicke et al., 2011; Olson et al., 2012b). Populations were found on several of the islands in the Philippines in residential areas with nursery and landscape plants (Olson et al., 2014; Sy et al., 2015a; Sy et al., 2015b; Sy and Salgo, 2015).

After initial introduction to novel sites via human mediated transport, E. planirostris then, in addition to human mediated transport, colonised additional sites unassisted (including disturbed and natural habitats) increasing its distribution in Florida, Louisiana, Hawaii, Guam and Alabama (Goin, 1944; Goin, 1947; Dundee, 1994; Butterfield et al., 1997; Johnson et al., 2003; Meshaka et al., 2009; Olson et al., 2012b, Alix et al., 2014).

Introductions

Risk of Introduction

E. planirostris is one of the most successful amphibian invaders (Bomford et al., 2009). Given that their primary mode of transport through the nursery trade is active and globally expanding, further introductions and establishment are likely. Establishment success in invaded habitats and dispersal risk of this species is high. Due to its inconspicuous nature and the habitats it occupies, initial infestations are often overlooked until populations are already too large and widespread to control (Kraus and Campbell, 2002; Olson et al., 2012b). Although most introductions will be restricted to tropical areas (Rödder and Lötters, 2010), long-term residence in the Florida Keys may have allowed E. planirostris to evolve adaptations to colder climates and populations from Florida may be able to spread into temperate zones (Heinicke et al., 2011). 

Habitat

E. planirostris is a terrestrial or sub-fossorial species, and can occassionally be found up to 2 m off the ground (Duellman and Schwartz, 1958). They are often found under objects such as leaf litter, rocks, gravel, fallen branches, coconut husks, flower pots, human debris, or hidden inside low-growing bromeliads, caves, lava tubes, rock crevices, gopher tortoise burrows, irrigation boxes, burrowing into loose, moist soil and under loose bark (Goin 1947; Neill 1951; Stewart and Martin 1980; Lips 1991; Schwartz and Henderson, 1991; Díaz and Cádiz, 2008; Olson and Beard 2012; Villa, 2015).

In Cuba, E. planirostris is found in wet and dry forests, coastal and mountainous areas, riversides, streambeds, caves, rocky outcrops, gardens and houses (Garrido and Schwartz, 1968; Díaz and Cádiz, 2008).

In its introduced range, it is often found in gardens, urban and residential areas and disturbed habitats, but will also be found in a variety of natural and forested areas. For example, in Florida, it is found in agricultural areas, disturbed areas, urban parks and gardens, mangrove forests, tropical hardwood hammocks, oak-palm hammocks, mesic hammocks, rockland hammocks, pine rockland, pine flatwoods, xeric uplands, scrub habitats, wet and dry prairie, freshwater marshes and coastal scrub areas (Meshaka, 2011). In Jamaica, it is found in open grasslands, scrub, lawns, pastures, roadsides and other disturbed areas (Stewart and Martin, 1980). In Panama and Mexico, records of E. planirostris are from urban areas, associated with houses and gardens and in grassy and forested areas close to residential sites (Crawford et al., 2011; Cedeño-Vázquez et al., 2014). In Hawaii, they are found in disturbed sites such as nurseries, roadsides, residential gardens, orchards and areas with native shrublands and forests (Olson et al., 2012a). In Guam, they are found in residential areas and secondary scrub forests (Olson et al., 2012b). Records from the Philippines are from urban areas with landscaped plants (Olson et al., 2014; Sy et al., 2015a; Sy et al., 2015b; Sy and Salgo, 2015).

Habitat List

Biology and Ecology

Genetics

Nucleotide sequencing of the mitochondrial cyt-b gene and nuclear rag-1 and pomc genes of E. planirostris indicates two separate lineages; an eastern and western lineage. The “eastern” lineage is found in eastern/central Cuba, the Bahamas and the Cayman Islands and has been introduced into Jamaica, Nicaragua (Miskito Cay) and North Caicos. The “western” lineage is found in western Cuba and has been introduced into Florida and Hawaii. Genetic variation is higher for eastern lineages and Cuba than for the introduced population from Florida (Heinicke et al., 2011). Genetic analyses of mitochondrial Cytochrome Oxidase Subunit I (COI) and a fragment of the ribosomal 16S gene confirmed species introductions to Panama City, Panama and Yucatán, Mexico (Crawford et al., 2011; Cedeño-Vázquez and others 2014). The Panama population are of the “western” lineage and may be a secondary introduction from Florida origin (Crawford et al., 2011).

Reproductive Biology

E. planirostris is a direct-development frog, which means there is no tadpole phase and complete metamorphosis takes place inside of the egg. Eggs are laid on the ground, or under fallen leaves, rocks, or other debris (Goin, 1947; Díaz and Cádiz, 2008). Clutch size ranges from 3-26 eggs, and eggs hatch 13-20 days after deposition (Goin, 1947). Hatchlings resemble adult frogs, with similar colour patterns and an egg-tooth and reduced tail that is absorbed (Goin, 1947). Snout-vent-length (SVL) ranges from 4.3-5.7 mm (Goin, 1947; Lazell, 1989).

In Cuba, vitellogenesis is bimodal with first occurrence from April to June and then again from September to October. Temperature is a vital environmental cue for onset. Spermatogenesis occurs from March to November peaking in June and July (Iturriaga et al., 2014). A description of the gonads is provided by Iturriaga et al. (2012).

E. planirostris breeds seasonally in Cuba from April through January (Meshaka and Layne, 2002) and April to September in Florida (Goin, 1947; Meshaka and Layne, 2002). In Louisiana, breeding is limited to June and July (Dundee and Rossman, 1989). Breeding is seasonal in Hawaii and mirror the breeding season of Florida (Ferreira et al., 2015). Breeding usually occurs at night in warm and humid conditions (Meshaka and Layne, 2002). Males will call on the ground or vegetation, under debris or rocks and from subterranean lava tubes (Díaz and Cádiz 2008; Olson et al., 2012b).

Frogs become reproductively active 6-8 months after hatching in south-central Florida (Meshaka and Layne, 2002) and after one year in north Florida (Goin, 1947). The mean survivorship in Florida is estimated to be 1.9 ± 2.3 months (range: 0.03-6.6) (Meshaka and Layne, 2002).

Activity Patterns

E. planirostris is active all year round in Cuba but in Florida, it may limit its activity during cooler temperatures (Goin, 1947). In the Everglades of southern Florida, E. planirostris was found hibernating under the bark of a wild tamarind tree (Harper, 1935). In native and introduced habitats, they are predominantly nocturnal, actively foraging at night and seeking daytime retreat sites in leaf litter or under debris (Goin, 1947; Stewart 1977; Díaz and Cádiz, 2008; Meshaka, 2011; Olson and Beard, 2012). Advertisement calls are heard at night with peak chorus just after sunset and before sunrise (Goin, 1947).

Population Size and Structure

Population density estimates are only available from introduced habitats. Densities estimated from mark-recapture studies of adults at three sites on the island of Hawaii were 4,564, 2,400 and 5,300 adults ha⁻¹ (Olson et al., 2012a; Olson et al., 2012b). Preadult to adult ratio was also determined at the first site as 1.7, thus the total population density was estimated to be 12,522 frogs ha^-1. In Jamaica, the density of E. planirostris and three other Eleutherodactylus species was estimated at 4,635 frogs ha^-1. The abundance of E. planirostris was found to be higher in the dry season, at coastal regions and sites without native species (Stewart and Martin, 1980). For the latter survey, it was the most abundant species found. In southeastern Florida, it was one of the most abundant species during a study using coverboards (Engeman et al., 2016), In southwestern Florida, it was the only anuran species, native or exotic, in which mean calling intensity increased over a ten year period (Everham et al., 2013).

Nutrition

Dietary studies indicate that E. planirostris is primarily an insectivore and predominantly consumes leaf litter invertebrates. They will eat a variety of available species, but may specialise on ants (Olson and Beard, 2012; Ferreira et al., 2015). In Cuba, the native range, ants comprised 100% of their diet (Goin, 1947). In its introduced habitats of Florida, Jamaica and Hawaii, ants were also the dominant prey (Goin, 1947; Stewart, 1977; Olson and Beard, 2012; Ferreira et al., 2012). Other invertebrates found in their diet include springtails, arachnids (spiders and mites), termites, beetles, collembola, amphipods, isopods, cockroaches, dermaptera, millipedes, centipedes, worms, flies (adults and larvae), hemiptera (true bugs), Lepidoptera larvae and snails.

Associations

In southeastern USA, E. planirostris is one of many species that are found commensal with the gopher tortoise (Gopherus polyphemus) using its burrows in mesic and xeric forests (Lips, 1991; Witz et al., 1991). One study in a long-leaf pine savannah habitat indicated that there is a slight correlation between E. planirostris abundance and number of gopher tortoise burrows (Catano and Stout, 2015).

Environmental Requirements

E. planirostris has a high tolerance for warm and dry conditions compared to other Jamaican Eleutherodactylus species, with a preferred temperature of 27.3 ± 0.66°C (Pough et al., 1977). The maximum temperature range for E. planirostris was found to be from 36.4 to 41.8°C and critical water loss at 34.9% ± 0.004 of initial body weight in 40-50% relative humidity (Pough et al., 1977). Most areas of introductions have similar mean annual and maximum warmest-month temperatures to Cuba (Rödder and Lötters, 2010). Temperature and rainfall will likely limit its spread in temperate climates; for example, populations found in Oklahoma City and Missouri are limited to indoor greenhouses and buildings (US Geological Survey, 2016). However, there is a record of one individual that survived one month in a frozen mulch bag in Michigan (Zippel et al., 2005) and given their long term residence in peninsular Florida, E. planirostris may have evolved adaptations to survive colder climates (Bomford et al., 2009; Heinicke et al., 2011). Species distribution models indicate that E. planirostris exhibits niche conservatism in its invaded habitats but this may be due to the environmental gradients available (Rödder and Lötters, 2010). It is believed that the distribution of E. planirostris is increasing northward as a result global warming.

E. planirostris seems to be restricted to low elevation areas. They are found from sea level up to 720 m in Cuba, 600 m in Jamaica and 500 m in the southeastern USA and Hawaii (Stewart and Marin, 1980; Díaz and Cádiz, 2008; Olson et al., 2012b).

Climate

Latitude/Altitude Ranges

Natural enemies

Notes on Natural Enemies

Snakes are a predator of E. planirostris in its native and introduced range, including three species of native racers; Cubophis canterigerus in Cuba, C. caymanus on Grand Cayman and C. vudii in the Bahamas (Henderson and Powell, 2009). In Florida, the native ringneck snake, Diadophis punctatus, a small, fossorial species found in humid and moist habitats, is a predator of E. planirostris (Wilson and Porras, 1983; Lazell, 1989). In Guam the invasive brown tree snake, Boiga irregularis, has been documented consuming E. planirostris (Mathies et al., 2012).

Other species of frog are also predators of E. planirostris. On Grand Cayman, E. planirostris was found in stomach content analyses of the Cuban treefrog, Osteopilus septentrionalis (Meshaka, 1996). Other predators in its native and introduced range probably also include invertebrates, lizards, birds and mammals (Henderson and Powell, 1999).

Documented parasites in its native Cuba include the nematodes Batracholandros bassi and Oswaldocruzia lenteixeirai (Goldberg and Bursey, 1991; Henderson and Powell 2009). Intestinal fluke parasites from the Mesocoeliidae family, Mesocoelium meggitti and M. monodi were found in introduced E. planirostris from the southeastern USA (Dronen et al., 2012). E. planirostris has also tested positive for the chytrid fungus, Batrachochytrium dendrobatidis, in its introduced range in Florida and Jamaica (Rizkalla, 2010; Holmes et al., 2012).

Means of Movement and Dispersal

Natural Dispersal

Dispersal of E. planirostris via oceanic crossings is possible. Self-propelled natural dispersal is thought to be the mode of introduction of this species into the Bahamas, the Cayman Islands and the Florida keys (Goin, 1944; Goin, 1947; Heinicke et al., 2007; Meshaka et al., 2009; Heinicke et al., 2011; Meshaka, 2011).

Accidental Introduction

E. planirostris is a terrestrial species that lays its eggs in moist soil or plants. The adults and juveniles are small and dark-coloured and the call is inconspicuous. As a result, they are transported, unintentionally and undetected, as hitchhikers in cargo, mostly in shipments of nursery plants and materials (Christy et al., 2007a; Kraus, 2009; Olson et al., 2012a). They thrive in plant nurseries and landscaped, residential areas and adults, juveniles and eggs are all transported in soil and mulch, potted plants, or leaf rosettes (Zippel et al., 2005; Christy et al., 2007a). Accidental introduction is the only method of E. planirostris transport to the Caribbean and southeastern USA, outside of the Florida Keys, Guam and the Philippines via the cargo or the horticulture trade (Stewart, 1977; Wilson and Porras, 1983; Dundee et al., 1989; Platt and Fontenot, 1995; Kraus et al., 1999; Winn et al., 1999; Dinsmore, 2004; Zippel et al., 2005; Christy et al., 2007a; Kraus, 2009; Meshaka et al., 2009; Olson et al., 2014; US Geological Survey, 2016).

Impact Summary

Economic Impact

In Hawaii, where high densities of E. planirostris have been reported around resorts and hotels, costs have been incurred to treat infestations of irrigation boxes and around swimming pools however, these are minimal (Olson et al., 2012b).

Environmental Impact

E. planirostris consumes a variety of leaf-litter invertebrates which could threaten native species of ants, mites, spiders and beetles (Goin, 1947; Duellman and Schwartz, 1958; Olson and Beard, 2012). It is possible that this species may compete with native insectivores, such as the Reef Gecko (Sphaerodactylus notatus) in southern Florida (Meshaka, 2011). Additionally, changes in native leaf litter invertebrate communities can impact ecosystem processes such as nutrient cycling or leaf litter decomposition rates, however as yet, there is no data to prove this (Olson and Beard, 2012). In Guam, E. planirostris may also provide an unnatural prey source for the invasive brown tree snake, Boiga irregularis and facilitate its spread (Mathies et al., 2012).

In Jamaica, E. planirostris have tested positive for the presence of Batrachochytrium dendrobatidis, the amphibian chytrid fungus (Holmes et al., 2012) which may be a vector risk to native amphibians.

Nevertheless, in Hawaii, where all ant species are non-native and many negatively impact native invertebrates, plants and vertebrates, it is possible that E. planirostris may have a positive impact as it may help reduce non-native ant populations, specifically big-headed ants, Pheidole megacephala, Argentine ants, Linepithema humile and yellow crazy ants, Anoplolepis gracilipes (Olson and Beard, 2012).

Social Impact

The social impact of E. planirostris has not been studied but large populations can be a nuisance to private home owners and property managers (Kraus et al., 1999; Kraus and Campbell, 2002).

Risk and Impact Factors

• Proved invasive outside its native range
• Has a broad native range
• Abundant in its native range
• Is a habitat generalist
• Tolerant of shade
• Capable of securing and ingesting a wide range of food
• Highly mobile locally
• Benefits from human association (i.e. it is a human commensal)
• Has high reproductive potential

• Changed gene pool/ selective loss of genotypes
• Negatively impacts agriculture
• Negatively impacts livelihoods
• Negatively impacts tourism
• Reduced amenity values
• Transportation disruption
• Negatively impacts animal/plant collections
• Negatively impacts trade/international relations

• Competition - monopolizing resources
• Interaction with other invasive species
• Predation
• Rapid growth

• Highly likely to be transported internationally accidentally
• Difficult to identify/detect as a commodity contaminant
• Difficult to identify/detect in the field
• Difficult/costly to control

Diagnosis

Genetic analyses involving sequencing of mitochondrial DNA was conducted to confirm the introduction of E. planirostris into Panama City, Panama, Yucatán, Mexico and Isla de Guanaja, Honduras (Crawford et al, 2011; Cedeño-Vázquez et al., 2014; McCranie et al., 2014). The results from Panama were deposited at the Barcode of Life Data Systems (Ratnasingham and Hebert, 2007) under project code “BSINV” and from Mexico Caribbean, under project code “EMC.”

Detection and Inspection

E. planirostris is fairly quiet and inconspicuous and may be difficult to detect until introduced populations are too large to manage (Olson et al., 2012a). In Hawaii, visual and aural inspections of plant shipments may occur but are somewhat ineffective due to the small size of the frogs and eggs and because inspections are conducted during the day, when males are not usually actively calling (Olson et al., 2012b). A number of keys and descriptions are provided to enable accurate identification (Conant and Collins, 1998; Díaz and Cádiz, 2008; Köhler, 2011). 

Similarities to Other Species/Conditions

E. planirostris is similar in appearance to a number of species within the same genus. For example, in Cuba, both E. ricordii and E. goini are similar but are larger than E. planirostris (Schwartz 1965; Schwartz, 1974). E. casparii is also similar but has black bands on the sides of the body behind the front limbs and a greenish tint to the dorsal colouring. E. tonyi and E. simulans, are almost identical to E. planirostris but can be differentiated by advertisement calls (Díaz and Cádiz, 2008). These species however, have not been introduced into areas outside of their native range.

In the southeastern USA, its introduced range, Acris crepitans and A. gryllus, are distinguished by their warty skin and dark stripes on the inside of their thighs and black triangle under their eyes and spring peepers and Pseudacris crucifer, is distinguished by an "X" mark on its back (Bartlett and Bartlett, 1999). These species also have very different advertisement calls.

In Hawaii, the invasive and closely related species E. coqui is distinguished by its larger size and toe pads, wider snout and loud, two-note advertisement call (Olson et al., 2012b). In the Philippines, some terrestrial species of Platymantis superficially resemble E. planirostris, but their calls are distinctly different (Olson et al., 2014).

Prevention and Control

Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Prevention

Early Warning Systems/Rapid Response

The US Geological Society (USGS) has an online sighting report form where new locations of Nonindigenous Aquatic Species (NAS) can be reported for the country (US Geological Survey, 2016).

Public Awareness

Public awareness is required to prevent the spread of E. planirostris from nurseries and, where possible to help control local populations. A series of newspaper articles have been published detailing the threats of invasive E.coqui and E. planirostris in Hawaii (Thompson, 2000; Thompson, 2001; Honolulu Star-Bulletin, 2001; Fujimori, 2001; Chun et al., 2003; Raloff, 2003; CTAHR, 2016; HEAR, 2016). These were however produced too late to control these populations.

Control

Cultural Control and Sanitary Measures

Leaf litter and debris removal may reduce the abundance of E. planirostris in privately owned areas (Olson et al., 2012b; CTAHR, 2016).

Physical/Mechanical Control

Hand capture may not be an efficient control method for the cryptic and elusive E. planirostris because abundances may actually be quite large once an introduction is detected and the frog is too quick to capture (Olson et al., 2012b). Native habitat management of fire-adapted communities may be an effective means of control in the southeastern USA (Meshaka, 2011).

Movement Control

Traps and barriers that were developed in Hawaii for E. coqui may work on E. planirostris and could be placed at invasion fronts or to prevent spread to vulnerable biodiverse habitats or privately owned areas. Hot water spray and vapour treatments of plant shipments has been effective for controlling E. coqui and may be employed by plant nurseries, quarantines zones and residential areas to prevent the spread of E. planirostris (Olson et al., 2012b).

Chemical Control

In Hawaii, a variety of chemical substances has proved effective in reducing populations of Eleutherodactylus species. Most applications have however specifically targeted E. coqui. Citric acid is the only approved chemical for controlling species of Eleutherodactylus in Hawaii and has been shown to be effective on E. planirostris (Pitt and Sin, 2004; Olson et al., 2012b; CTAHR, 2016).

Gaps in Knowledge/Research Needs

With new records indicating an expanding global population it is important that further studies on this species are undertaken (Rödder and Lötters, 2010). Such studies should focus on the biology of E. planirostris in both native and introduced habitats; conducting studies to determine minimum temperature tolerances are needed. This would help to understand the risk of E. planirostris spreading to temperate climates.

In most of its introduced range, it is unknown if E. planirostris is invasive and impacting native ecosystems. Comprehensive diet studies, including evidence of prey preference and demographic impacts on native species, should be conducted in invaded habitats outside of Hawaii (Olson and Beard, 2012; Ferreira et al., 2015). Although it may compete with native species in many of its introduced habitats, no specific studies have studied competition with native species. Large densities E. planirostris may be altering its prey (invertebrate) communities and impacts to ecosystem processes should be assessed. Studies should also examine the effect that large densities of E. planirostris have on both native and non-native predator populations (Mathies et al., 2012; Cassani et al., 2015). 

References

Alix DM; Guyer C; Anderson CJ, 2014. Expansion of the range of the introduced Greenhouse Frog, Eleutherodactylus planirostris, in coastal Alabama. Southeastern Naturalist, 13(4):N59-N62. http://www.bioone.org/doi/full/10.1656/058.013.0409

Ashton RE; Ashton PS, 1988. Handbook of reptiles and amphibians of Florida: part three: the amphibians. Miami, Florida, USA: Windward Publishing, 191 pp.

Barbour T, 1910. Eleutherodactylus ricordii in Florida. Proceedings of the Biological Society of Washington, 23:100.

Barbour T; Ramsden CT, 1919. The herpetology of Cuba. Memoirs of the Museum of Comparative Zoology at Harvard College, 47(2):71-213.

Bartlett RD; Bartlett P, 2006. Guide and reference to the amphibians of eastern and central North America. Gainesville, Florida, USA: University Press Florida, 280 pp.

Bomford M; Kraus F; Barry SC; Lawrence E, 2009. Predicting establishment success for alien reptiles and amphibians: a role for climate matching. Biological Invasions, 11(3):713-724. http://www.springerlink.com/content/bv005082j8m4j678/?p=02379768c67c4698b29a060f692252be&pi=20

Boundy J, 1998. Distributional records for Louisiana amphibians. Herpetological Review, 29(4):251-252.

Boundy J, 2004. Amphibian and reptile distribution records for Louisiana. Herpetological Review, 35(2):194-196.

Breuil M, 2002. [English title not available]. (Histoire naturelle des amphibiens et reptiles terrestres de l'archipel guadeloupéen (Guadeloupe, Saint-Martin, Saint-Barthélémy: Basse-Terre, Grande-Terre et les îlets satellites, Marie-Galante, les Saintes, la Désirade, les îles de la Petite Terre, Saint-Martin et les îlets satellites, Saint-Barthélémy et îlets satellites).) MNHN, Institut d'Écologie et de Gestion de la Biodiversité, Service du Patrimoine Naturel, Patrimoines naturels No. 54. Paris, France.

Butterfield BP; Meshaka Jr WE; Guyer C, 1997. Nonindigenous amphibians and reptiles, strangers in paradise: impact and management of non-indigenous species in Florida. Island Press. Washington D.C., USA 123-137.

Carey SD, 1982. Geographic distribution: Eleutherodactylus planirostris. Herpetological Review, 13(4):130.

Cassani JR; Croshaw DA; Bozzo J; Brooks B; Everham EM III; Ceilley DW; Hanson D, 2015. Herpetofaunal community change in multiple habitats after fifteen years in a Southwest Florida Preserve, USA. PLoS ONE, 10(5):e0125845. http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0125845

Catano CP; Stout IJ, 2015. Functional relationships reveal keystone effects of the gopher tortoise on vertebrate diversity in a longleaf pine savanna. Biodiversity and Conservation, 24(8):1957-1974. http://link.springer.com/article/10.1007%2Fs10531-015-0920-x

Cedeño-Vázquez JR; González-Vázquez J; Martínez-Arce A; Canseco-Márquez L, 2014. First record of the invasive greenhouse frog (Eleutherodactylus planirostris) in the Mexican Caribbean. Revista Mexicana de Biodiversidad, 85(2):650-653. http://www.ib.unam.mx/revista/

Christy MT; Clark CS; Gee DE; Vice D; Vice DS; Warner MP; Tyrrell CL; Rodda GH; Savidge FA, 2007. Recent records of alien anurans on the Pacific Island of Guam. Pacific Science, 61:469-483.

Christy MT; Savidge JA; Rodda GH, 2007. Multiple pathways for invasion of anurans on a Pacific island. Diversity and Distributions, 13(5):598-607. http://www.blackwell-synergy.com/loi/ddi

Chun S; Hara AH; Niino-DuPonte RY, 2003. Greenhouse frog or coqui? Differences between two Eleutherodactylus species found in Hawaii. Manoa, Hawaii, USA: College of Tropical Agriculture and Human Resources (CTAHR), University of Hawaii.

Conant R; Collins JT, 1998. A field guide to reptiles and amphibians: eastern and central North America. Third Edition, Expanded. Boston, Massachusetts, USA: Houghton Mifflin Company, 616 pp.

Cope ED, 1863. On Trachycephalus, Scaphiopus, and other American Batrachia. Proceedings of the Academy of Natural Sciences of Philadelphia, 15:43-54.

Cope ED, 1875. Check-list of North American Batrachia and Reptilia: with a systematic list of the higher groups, and an essay on geographical distribution. Bulletin of the United States National Museum, 1:i-xi,1-104.

Cope, 1889. The Batrachia of North America. Bulletin of the United States National Museum, 34:1-525.

Crawford AJ; Alonso R; Jaramillo A CA; Sucre S; Ibáñez D R, 2011. DNA barcoding identifies a third invasive species of Eleutherodactylus (Anura: Eleutherodactylidae) in Panama City, Panama. Zootaxa, 2890:65-67. http://www.mapress.com/zootaxa/2011/f/z02890p067f.pdf

Crombie RI, 1999. Jamaica. In: Caribbean amphibians and reptiles [ed. by Crother, B. I.]. San Diego, California, USA: Academic Press, 63-92.

CTAHR, 2016. Control of coqui frogs in Hawaii. Manoa, Hawaii, USA: College of Tropical Agriculture and Human Resources (CTAHR), University of Hawaii. http://www.ctahr.hawaii.edu/coqui/control.asp

Díaz LM; Cádiz A, 2008. [English title not available]. (Guía Taxonómica de los Anfibios de Cuba.) Abc taxa, 4. 294 pp.

Dillman JW; Gibbons PW, 2016. Geographic distribution: Eleutherodactylus planirostris (greenhouse frog). Herpetological Review, 47(1):76-77.

Dinsmore A, 2004. Geographic distribution: Eleutherodactylus planirostris (greenhouse frog). Herpetological Review, 35:403.

Dronen NO; Calhoun DM; Simcik SR, 2012. Mesocoelium Odhner, 1901 (Digenea: Mesocoelidae) revisited; a revision of the family and re-evaluation of species composition in the genus. Zootaxa, 3387:1-96.

Duellman WE; Schwartz A, 1958. Amphibians and reptiles of southern Florida. Bulletin of the Florida State Museum,Biological Sciences, 3:181-324.

Dundee HA, 1994. Geographical distribution: Eleutherodactylus planirostris planirostris (greenhouse frog). Herpetological Review, 25(4):160.

Dundee HA; Rossman DA; Beckham EC, 1989. The amphibians and reptiles of Louisiana. Baton Rouge, Louisiana, USA: Louisiana State University Press, 330 pp.

Echternacht AC; Burton FJ; Blumenthal JM, 2011. The amphibians and reptiles of the Cayman Islands: conservation issues in the face of invasions. In: Conservation of Caribbean Island herpetofaunas, vol 2 [ed. by Hailey, A. \Wilson, B. S. \Horrocks, J. A.]. Leiden, Netherlands: Brill., 129-147.

Elbers JP, 2007. Geographic distribution: Eleutherodactylus planirostris (greenhouse frog). Herpetological Review, 38(4):474.

Engeman RM; Meshaka WE Jr; Severson R; Severson MA; Kaufman G; Groninger NP; Smith HT, 2016. Monitoring cryptic amphibians and reptiles in a Florida state park. Environmental Science and Pollution Research, 23(7):7032-7037. http://link.springer.com/article/10.1007%2Fs11356-015-6028-8

Estrada A; R; Ruibal R, 1999. A review of Cuban herpetology. In: Caribbean amphibians and reptiles [ed. by Brian, I. C.]. San Diego, California, USA: Academic Press,, 31-62.

Everham IIIEM; Ceilley DW; Croshaw DA; Firth J; Gunnels C; Hanson DD; Mariolan S; Spear RJ; Thomas B; Norman DEVan; Whitmore BM; Cassani JR, 2013. Ten years of the southwest Florida frog monitoring network: natural variation and human-driven changes. Florida Scientist, 76(2):138-149.

Ferreira RB; Beard KH; Choi RT; Pitt WC, 2015. Diet of the nonnative greenhouse frog (Eleutherodactylus planirostris) in Maui, Hawaii. Journal of Herpetology, 49(4):586-593.

Frost DR; Grant T; Faivovich J; Bain RH; Haas A; Haddad CFB; Sà RO De; Channing A; Wilkinson M; Donellan SC; Raxworthy CJ; Campbell JA; Blotto BL; Moler P; Drewes RC; Nussbaum RA; Lynch JD; Green DM; Wheeler WC, 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History, 297:1-291.

Fujimori L, 2001. Shrieking frogs invade Oahu. Honolulu, Hawaii, USA: Honolulu Star-Bulletin. http://archives.starbulletin.com/2001/07/08/news/story1.html

García-Balderas CM; Cedeño-Vázquez JR; Mineros-Ramírez R, 2016. Distribution note: Eleutherodactylus planirostris (Cope, 1862). Mesoamerican Herpetology, 3(1):173.

Garrido OH; Schwartz A, 1968. [English title not available]. (Anfibios, reptiles y aves de la península de Guanahacabibes.) Poeyana, 53:1-68.

Godwin C, 2014. Geographic distribution: Eleutherodactylus planirostris (greenhouse frog). Herpetological Review, 45(3):458.

Goin CJ, 1944. Eleutherodactylus ricordii at Jacksonville, Florida. Copeia, 1944(3):192.

Goin CJ, 1947. Studies on the life history of Eleutherodactylus ricordii planirostris (Cope, 1862) in Florida: with special reference to the local distribution of an allelomorphic color pattern. Gainesville, Florida, USA: University of Florida Press, 66 pp.

Goin CJ, 1955. Description of a new subspecies of the frog Eleutherodactylus ricordi from the Bahama Islands. American Museum Novitates, 1708:1-7.

Goldberg SR; Bursey CR, 1996. Gastrointestinal helminths of the anole Anolis oculatus (Polychridae) from Dominica, Lesser Antilles. Journal of the Helminthological Society of Washington, 63(1):125-128.

González VR; Schettino LR; Mancina CA; Iturriaga M, 2014. Amphibians of Cuba: checklist and geographic distributions. Division of Amphibians and Reptiles, National Museum of Natural History, Smithsonian Institution, 145:1-48.

Graham SP; Timpe EK; Giovanetto; L, 2007. Significant new records for Georgia herpetofauna. Herpetological Review, 38(4):494-496.

Grant C, 1940. The herpetology of the Cayman Islands. Bulletin of the Institute of Jamaica, Science Series, 2:1-65.

Greene M, 2014. Geographical distribution: Eleutherodactylus planirostris (greenhouse frog). Herpetological Review, 44(4):620.

Groenewoud D; Law IS, 2016. First record of greenhouse frog in Singapore. Singapore Biodiversity Records. http://lkcnhm.nus.edu.sg/nus/images/pdfs/sbr/2016/sbr2016-059.pdf

Harper F, 1935. Records of amphibians in the southeastern states. The American Midland Naturalist, 16(3):275-310.

HEAR, 2016. Stop the spread of Caribbean frogs. Pest Alert!. Hawaii, USA: Hawaii Ecosystems at Risk (HEAR). http://www.hear.org/alienspeciesinhawaii/species/frogs/frogalert20000404.pdf

Hedges SB, 1999. Distribution patterns of amphibians in the West Indies. In: Patterns of distribution of amphibians: a global perspective [ed. by Duellman, W.]. Baltimore, Maryland, USA: The Johns Hopkins University Press, 211-254.

Hedges SB; Duellman WE; Heinicke MP, 2008. New World direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737:1-182.

Heinicke MP; Diaz LM; Hedges SB, 2011. Origin of invasive Florida frogs traced to Cuba. Biology Letters, 7(3):407-410. http://rsbl.royalsocietypublishing.org/content/7/3/407.full

Heinicke MP; Duellman WE; Hedges SB, 2007. Major Caribbean and Central American frog faunas originated by ancient oceanic dispersal. Proceedings of the National Academy of Sciences of the United States of America, 104:10092-10097.

Henderson RW; Powell R, 1999. West Indian herpetoecology. In: Caribbean amphibian and reptiles [ed. by Crother, B. I.]. San Diego, California, USA: Academic Press, 223-268.

Henderson RW; Powell R, 2009. Natural history of West Indian reptiles and amphibians. University Press of Florida. Gainesville, Florida, USA, 495 pp.

Holmes I; McLaren K; Wilson B, 2012. Surveys for frog diversity and Batrachochytrium dendrobatidis in Jamaica. Herpetological Review, 43(2):278.

Honk Kong Bird Watching Society (HKBWS), 2013. Hong Kong headline indicators for biodiversity and conservation - 2011 report., Hong Kong: Hong Kong Bird Watching Society, 26 pp. http://www.hkbws.org.hk/web/chi/documents/2012_HK_Headline_Indicators_for_Biodiversity_Conservation.pdf

Honolulu Star-Bulletin, 2001. Tiny, alien frogs making big noise across Hawaii. Honolulu, Hawaii, USA: Honolulu Star-Bulletin. http://archives.starbulletin.com/2001/05/18/editorial/editorials.html

Hyning OC van, 1933. Batrachia and Reptilia of Alachua County, Florida. Copeia, 1933(1):3-7.

iNaturalist, 2016. iNaturalist. California, USA: University of California Berkley. http://www.inaturalist.org/

Iturriaga M; Rodríguez Y; Sanz A, 2012. Structural and ultrastructural description of the gonads of Eleutherodactylus planirostris (Anura: Eleutherodactylidae). Herpetology Notes, 5:281-290.

Iturriaga M; Sanz A; Oliva R, 2014. Seasonal Reproduction of the greenhouse frog Eleutherodactylus planirostris (Anura: Eleutherodactylidae) in Havana, Cuba. South American Journal of Herpetology, 9(2):142-150.

Jensen JB; Camp CD; Gibbons JW; Elliott MJ, 2008. Amphibians and reptiles of Georgia. Athens, Georgia, USA: University of Georgia Press, 575 pp.

Johnson SA; Staiger JS; Barichivich WJ, 2003. Geographic distribution: Eleutherodactylus planirostris (greenhouse frog). Herpetological Review, 34(2):161-162.

Kaiser H, 1992. The trade-mediated introduction of Eleutherodactylus martinicensis (Anura: Leptodactylidae) on St. Barthélémy, French Antilles, and its implications for Lesser Antillean biogeography. Journal of Herpetology, 26(3):264-273.

Kraus F, 2009. Invading nature: Springer series in invasion ecology 4. Springer, 563 pp.

Kraus F; Campbell EW; Allison A; Pratt T, 1999. Eleutherodactylus frog introductions to Hawaii. Herpetological Review, 30:21-25.

Kraus F; Campbell EW; III, 2002. Human-mediated escalation of a formerly eradicable problem: the invasion of Caribbean frogs in the Hawaiian Islands. Biological Invasions, 4:327-332.

Krysko KL; Enge KM; Moler PE, 2011. Atlas of amphibians and reptiles in Florida. Final Report No. Project Agreement 08013,. Tallahassee, Florida, USA: lorida Fish and Wildlife Conservation Commission, 524 pp. http://www.flmnh.ufl.edu/herpetology/florida-amphibians-reptiles/checklist-atlas

Köhler G, 2011. Amphibians of Central America. Offenbach, Germany: Herpeton, Verlag Elke Köhler, 379 pp, 379 pp.

Lamb JY; Chatfield MWH, 2014. Geographic distribution: Eleutherodactylus planirostris (greenhouse frog). Herpetological Review, 45:458.

Lazell JD, 1989. Wildlife of the Florida Keys: a natural history. Washington D.C, USA: Island Press, 250 pp, 250 pp.

Lever C, 2003. Naturalized reptiles and amphibians of the world. Oxford, UK: Oxford University Press, 318 pp.

Liner E, 2007. Geographic distribution: Euhyas planirostris (greenhouse frog). Herpetological Review, 38(2):214.

Lips KR, 1991. Vertebrates associated with tortoise (Gopherus polyphemus) burrows in four habitats in south-central Florida. Journal of Herpetology, 25:447-481.

Lorvelec O; Pascal M; Pavis C; Feldmann P, 2007. Amphibians and reptiles of the French West Indies: iInventory, threats and conservation. Applied Herpetology, 4(2):131-161.

Lynn WG, 1937. Two new frogs from Jamaica. Herpetologica, 1(3):88-91.

Lynn WG, 1940. The Herpetology of Jamaica. Bulletin of the Institute of Jamaica,Science Series, 1:1-12.

Lynn WG; Dent JN, 1943. Notes on Jamaican amphibians. Copeia, 1943(4):234-242.

Lynn WG; Grant C, 1940. The herpetology of Jamaica. Bulletin of the Institute of Jamaica Science Series:1-148.

Mann DL; Mann T; Winstead N; Lu W, 2014. Geographical distribution: Eleutherodactylus planirostris (greenhouse frog). Herpetological Review, 45(4):652.

Mathies T; Pitt WC; Rabon JA, 2012. Boiga irregularis (brown treesnake): diet. Herpetological Review, 43(1):143-144.

McConnell R; McConnell T; Guyer C; Laurencio D, 2015. Geographical distribution: Eleutherodactylus planirostris (greenhouse frog). Herpetological Review, 46(4):559.

McCranie JR; Collart JR; Castaneda FE; Solis JM, 2008. Geographic distribution. Eleutherodactylus (Euhyas) planirostris (greenhouse frog). Herpetological Review, 39(3):362-363.

McCranie JR; Valdés Orellana L, 2014. New island records and updated nomenclature of amphibians and reptiles from the Islas de la Bahía, Honduras. Herpetology Notes, 7:41-49.

Meshaka Jr WE, 1996. Diet and colonization of buildings by the Cuban treefrog (Osteopilus septentrionalis) (Anura: Hylidae). Caribbean Journal of Science, 32:59-63.

Meshaka Jr WE; Jeff B; Avery AW, 2009. The dispersal of the greenhouse frog, Eleutherodactylus planirostris (Anura: Eleutherodactylidae), in Louisiana, with preliminary observations on several potential exotic colonizing species. Journal of Kansas Herpetology, 32:13-16.

Meshaka Jr WE; Layne J, 2002. Herpetofauna of a long-unburned sandhill habitat in south-central Florida. Florida Scientist, 65:35-50.

Meshaka WE Jr, 2011. A runaway train in the making: the exotic amphibians, reptiles, turtles, and crocodilians of Florida. Herpetological Conservation and Biology, 6(Monograph 1):iii + 94 pp. http://herpconbio.org/Volume_6/Monographs/Meshaka_2011.pdf

Meshaka WE Jr; Butterfield BP; Hauge JB, 2004. The exotic amphibians and reptiles of Florida [ed. by Meshaka, W. E.\Butterfield, B. P.\Hauge, J. B.]. Malabar, USA: Krieger Publishing Company, x + 155 pp.

Neill WT, 1951. Florida's air-plants and their inhabitants. Florida Naturalist, 24:61-66.

Olson CA; Beard KH, 2012. Diet of the introduced Greenhouse Frog in Hawaii. Copeia, No.1:121-129. http://www.bioone.org/doi/full/10.1643/CE-11-008

Olson CA; Beard KH; Koons DN; Pitt WC, 2012. Detection probabilities of two introduced frogs in Hawaii: implications for assessing non-native species distributions. Biological Invasions, 14(4):889-900. http://www.springerlink.com/content/f51874w08110283x/

Olson CA; Beard KH; Pitt WC, 2012. Biology and impacts of Pacific island invasive species. 8. Eleutherodactylus planirostris, the greenhouse frog (Anura: Eleutherodactylidae). Pacific Science, 66(3):255-270. http://www.uhpress.hawaii.edu/journals

Olson CA; Diesmos A; Beard KH, 2014. Geographical distribution: Eleutherodactylus planirostris (greenhouse frog). Herpetological Review, 45(4):652-653.

Ouboter PE; Jairam R, 2012. Amphibians of Suriname. Leiden, Netherlands: Koninklijke Brill NV, 388 oo.

Petrovic CA, 1973. An "albino" frog, Eleutherodactylus planirostris Cope. Journal of Herpetology, 7(1):49-51.

Pitt WC; Sin H, 2004. Dermal toxicity of citric acid based pesticides to introduced Eleutherodactylus frogs in Hawaii. QA992 Final Report. Hilo, Hawaii, USA: USDA/APHIS/WS/National Wildlife Research Center.

Platt SG; Fontenot LW, 1995. Geographical distribution: Eleutherodactylus planirostris (greenhouse frog). Herpetological Review, 26(4):207.

Plotkin M; Atkinson R, 1979. Geographical distribution: Eleutherodactylus planirostris (greenhouse frog). Herpetological Review, 10(2):59.

Po; MK; Hang LY; Wyman K, 2016. Campus frog database., Hong Kong: Tree, Amphibian, Bird, Butterfly. City University of Hong Kong. http://www.tab2cityu.org/content.php?id=7

Pough FH; Stewart MM; Thomas RG, 1977. Physiological basis of habitat partitioning in Jamaican Eleutherodactylus. Oecologia, 27:285.

Powell R; Henderson RW; Eds, 2012. Island lists of West Indian amphibians and reptiles. Bulletin of the Florida Museum of Natural History, 51(2):85-166.

Powell R; Henderson RW; Farmer MC; Breuil M; Echternacht AC; Buurt Gvan; Romagosa CM; Perry G, 2011. Introduced amphibians and reptiles in the greater Caribbean: patterns and conservation implications. In: Conservation of Caribbean Island herpetofaunas, vol 2 [ed. by Hailey, A. \Wilson, B. S. \Horrocks, J. A. \editor]. Leiden, The Netherlands: Brill, 63-143.

Raloff J, 2003. Hawaii's hated frogs: tiny invaders raise a big ruckus. Science News, 163:11-13.

Ratnasingham S; Hebert PDN, 2007. The barcode of life data system (http://www.barcodinglife.org). Molecular Ecology Notes, 7(3):355-364.

Reichard SM; Stevenson HM, 1964. Records of Eleutherodactylus ricordi at Tallahassee. Florida Naturalist, 37:97.

Reynolds RG, 2011. Status, conservation, and introduction of amphibians and reptiles in the Turks and Caicos Islands, British West Indies. In: Conservation of Caribbean Island herpetofaunas, vol 2 [ed. by Hailey, A. \Wilson, B. S. \Horrocks, J. A. \editor]. Leiden, The Netherlands: Brill, 377-406.

Reynolds RG, 2012. Distribution: Eleutherodactylus planirostris (Cuban flathead frog) 27:1. Caribbean Herpetology, 27:1. http://www.caribbeanherpetology.org/pdfs/ch27

Reynolds RG; Niemiller ML, 2010. Island invaders: invasive reptiles and amphibians in the Turks and Caicos Islands. Reptiles & Amphibians, 17:1116-121.

Rizkalla CE, 2010. Increasing detections of Batrachochytrium dendrobatidis in central Florida, USA. Herpetological Review, 41(2):180.

Rödder D; Lötters S, 2010. Explanative power of variables used in species distribution modelling: an issue of general model transferability or niche shift in the invasive Greenhouse frog (Eleutherodactylus planirostris). Naturwissenschaften, 97(9):781-796. http://www.springerlink.com/content/l305340876177312/

Schwartz A, 1965. Geographic variation in two species of Hispaniolan Eleutherodactylus with notes on Cuban members of the ricordi group. Studies on the Fauna of Curacao and Other Caribbean Islands, 86:98-123.

Schwartz A, 1974. Eleutherodactylus planirostris. Catalogue of American Amphibians and Reptiles, 154:1-4.

Schwartz A; Henderson RW, 1991. Amphibians and reptiles of the West Indies: descriptions, distributions, and natural history. Gainesville, Florida, USA: University Press of Florida, 720 pp.

Schwartz A; Thomas RM; Ober LD, 1975. A check-list of West Indian amphibians and reptiles and first supplement. Carnegie Museum of Natural History Special Publication, 1:1-216.

Seidel ME; Franz R, 1994. Amphibians and reptiles (exclusive of marine turtles) of the Cayman Islands. In: The Cayman Islands: natural history and biogeography [ed. by Brunt, M. A. \Davies, J.]., The Netherlands: Kluwer Academic Publishers, 407-433.

Solís JM; Lainez JD; Salmeron MB, 2014. Distribution note: Eleutherodactylus planirostris (Cope, 1862). Mesoamerican Herpetology, 1(2):292.

Solís JM; Wilson LD; Townsend JH, 2014. An updated list of the amphibians and reptiles of Honduras, with comments on their nomenclature. Mesoamerican Herpetology, 1:123-144.

Stejneger L, 1917. Cuban amphibians and reptiles collected for the United States National Museum from 1899 to 1902. Proceedings of the United States National Museum, 53:33.

Stewart MM, 1977. The role of introduced species in a Jamaican frog community. Actas del IV Simposium Internacional de Ecologica Tropical:113-146.

Stewart MM; Martin GE, 1980. Coconut husk-piles-a unique habitat for Jamaican terrestrial frogs. Biotropica, 12:107-116.

Sy EY; Clifton J; Diesmos A, 2015. Geographical distribution: Eleutherodactylus planirostris (greenhouse frog). Herpetological Review, 46(1):56.

Sy EY; Eleazar IIIB; Achacosco R; Diesmos A, 2015. Geographical distribution: Eleutherodactylus planirostris (greenhouse frog). Herpetological Review, 46(1):56.

Sy EY; Salgo J, 2015. Geographical distribution: Eleutherodactylus planirostris (greenhouse frog). Herpetological Review, 46(2):212.

Thawley CJ, 2012. Geographical distribution: Eleutherodactylus planirostris (greenhouse frog). Herpetological Review, 43(2):298.

Thompson R, 2000. Pesky alien species have folks hopping mad. Honolulu, Hawaii, USA: Honolulu Star-Bulletin. http://archives.starbulletin.com/2000/06/05/news/story2.html

Thompson R, 2001. Caffeine kills unwanted frogs but EPA worries it may be too potent. Honolulu, Hawaii, USA: Honolulu Star-Bulletin. http://archives.starbulletin.com/2001/05/16/news/story4.html

Tipton BL; Hibbitts TL; Hibbitts TD; Hibbitts TJ; LaDuc TJ, 2012. Texas amphibians: a field guide. Austin, Texas, USA: University of Texas Press, 330 pp.

Ukpong EE; Jacob DE; Ufot IN; Joseph AE, 2012. The fauna biodiversity of Ikot Ondo Community forest in Essien Udim L. G. A. of Akwa Ibom State, South Eastern Nigeria. Ethiopian Journal of Environmental Studies and Management, 5(3):276-280. http://www.ajol.info/index.php/ejesm/article/view/77960

US Geological Survey, 2016. Nonindigenous Aquatic Species Database. Gainesville, USA: US Geological Survey. http://nas.er.usgs.gov

Villa J, 2015. [English title not available]. (Las ranitas de Cayos Miskitos.) Temas Nicaragüenses, 2015(89). 6-22.

Williams AA; Wygoda ML, 1997. Geographical distribution: Eleutherodactylus planirostris (greenhouse frog). Herpetological Review, 28(4):207.

Wilson BS, 2011. Conservation of Jamaica amphibians and reptiles. In: Conservation of Caribbean Island herpetofaunas vol 2 [ed. by Hailey, A. \Wilson, B. S. \Horrocks, J. A.]. Leiden, The Netherlands: Brill, 273-310.

Wilson LD; Porras L(, 1983. The ecological impact of man on the South Florida herpetofauna. Univ Kansas Mus Nat Hist Spec Publ, 9:1-171.

Winn B; Jensen JB; Johnson S, 1999. Geographical distribution: Eleutherodactylus planirostris (greenhouse frog). Herpetological Review, 30(1):49.

Witz BW; Wilson DS; Palmer MD, 1991. Distribution of Gopherus polyphemus and its vertebrate symbionts in three burrow categories. The American Midland Naturalist, 126(1):152-158.

Wright AH; Wright AA, 1949. Handbook of frogs and toads of the United States and Canada. 3rd edition. Ithaca, New York, USA: Comstock Publishing Company, Inc, 640 pp.

Zippel KC; Snider AT; Gaines L; Blanchard D, 2005. Eleutherodactylus planirostris (greenhouse frog): cold tolerance. Herpetological Review, 36:299-300.

Links to Websites

Contributors

17/05/2016 Original text by:

Christina Olson, Utah State University, USA

Distribution Maps