Incilius melanochlorus

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Incilius melanochlorus
Wet forest toad.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Amphibia
Order: Anura
Family: Bufonidae
Genus: Incilius
Species:
I. melanochlorus
Binomial name
Incilius melanochlorus
(Cope, 1877)
Synonyms

Bufo valliceps var.Cope, 1875 [2]
Bufo melanochlorusCope, 1877 [3]
Bufo melanochloris(Cope, 1877), lapsus [2] [3]
Cranopsis melanochlorus(Cope, 1877) [3]
Ollotis melanochlorus(Cope, 1877) [3]

Contents

Incilius melanochlorus, formerly Bufo melanochlorus, [4] [1] is a mid-sized species of toad with a crested head [2] in the family Bufonidae. [1] It is primarily distinguished by its very long first finger with respect to the other fingers. [2] It is found in southern Nicaragua, in the northern Cordillera Central (central highlands) and on the Atlantic slopes of eastern Costa Rica, and in western Panama. [3] [4]

Vernacular names

Frank and Ramus (1995) give it the common name dark green toad, [3] Panamanian herpetologists Jaramillo and Ibáñez (2009) use wet forest toad [4] or west forest toad according to Darrel R. Frost [3] (likely a misspelling). A local Spanish name from Costa Rica specifically for this species is sapo Costaricense de la selva, [5] but it also just known as sapo. [6]

Taxonomy

The first article about this species was published in 1875 by Edward Drinker Cope who wrote of the toad as a particular but unnamed variety of Bufo valliceps from eastern Costa Rica. It was described as a novel species in its own right in 1877 by Cope under the name B. melanochlorus. The holotype is a specimen (undesignated in Cope's original publication, but thought to be USNM30592) taken in Limón Province sometime before 1877 by W. W. Gabb. [2] [3] The specific epithet melanochlorus is compounded from the Ancient Greek μελανω (melanō), meaning "blackened" [7] and χλωρός (khlōrós) meaning “(yellowish) green”. [8] The English name invented by Frank and Ramus [3] is thus a simple transliteration.

In 2004 O'Neill and Mendelson moved this species from Bufo to Incilius , and while doing so split the taxon into two species, a western I. aucoinae and the nominate taxon. [1] Earlier publications use Bufo melanochlorus. [9] In 2006 Darrel Frost et al. moved the species to the old genus Cranopsis originally erected by Cope in 1875 with C. fastidiosus as the type species. [3] [10] It was controversial [11] and this approach was not followed by everyone. [1] [12] It subsequently necessitated moving Cranopsis toads to the new genus Cranophryne as the genus name Cranopsis proved to have already been used by Adams to describe a mollusc in 1860. [13] In 2009 Pauly et al. suggested that because the recent partitioning of Bufo had made the formerly monophyletic genus paraphyletic and that the new genera were too imperfectly defined to be phylogeneticly stable, as the large number of recent (officially mandated) name changes caused by Frost in 2006 was evidencing, and that the hitherto done genetic analyses were yet too vague to properly resolve relationships, thus Incilius should be treated as a subgenus of Bufo. [11] Frost refuted these criticisms in a 2009 response. A phylogenetic analysis by Mendelson based on morphology, life history, and molecular data was published 2011 which recommending sinking Cranopsis/Cranophryne, Ollotis and Crepidius/Crepidophryne back into Incilius. [14]

Mendelson et al. (2011) suggest that this species is part of an I. valliceps species group, or more specifically in what they termed the "Forest Group" subgroup, including the taxa I. aucoinae, I. cavifrons, I. campbelli, I. cristatus, I. leucomoyos, I. macrocristatus, I. spiculatus and I. tutelarius. [3]

Description

Incilius melanochlorus are moderately large toads, with males attaining 65 mm (2.6 in) [4] to 74 mm (2.9 in) [12] in snout–vent length, and females 103 mm (4.1 in) [4] to 107 mm (4.2 in). [12] There is marked sexual dimorphism, the females having more irregular dark colouration on their back and being larger, but the males having longer heads and more brawny forearms. [12]

The head is wider than it is long. [12] The head is crested, [2] with the cranium sporting well-developed, thin and high crests. [4] [12] This crest ornamentation consists of canthal, supraorbital, and postpreorbital crests, and parietal crests with transverse folds between them (other species have crests on other locations). The crests run from the nostril, behind the eye, to the back of the head. The skin of the top of the head is hardened and stuck to the top of the skull. [12] Males have a vocal sac. [4] The vocal slits which hide the vocal sac inside the throat are small and bilateral, the sac has a single lobe, and is large, heavily pigmented [12] with a greenish cast, [4] and when calling inflates fully to a round shape. The eyes are large with a coppery bronze iris and a black pupil, [12] the top half of the iris lighter than the bottom half, with a thin, dark line separating the two halves. [4] The tongue is long and thin. [12] The paratoid glands are small, [4] [12] smaller than the upper eyelid, [12] and triangular in shape, [4] [12] and the toad has a small tympanum. [12]

The hind legs are relatively short, as are the feet and the tibia, but the toes are long and have some webbing between them. The first finger is longer than the second. The fingers and toes have tubercles underneath them. Sometimes the tips of the fingers and toes are lighter-coloured, sometimes the same colour as the rest of the digit. The tarsus (heel) has two differently-shaped tubercles, one facing inwards, the other outwards. [12]

The dorsum (back) is very warty, [4] but is covered in a smooth skin except near the shoulders and hips, which has small, low and rounded protrusions. The ventral side has a minutely roughened surface. [12] The lateral side of the toad is bordered along the upper part with a row of light-coloured, low to medium-sized, sharp but rounded, spine-shaped warts extending from the paratoid gland to the groin. [4] [12] The upper surface of the hind limbs is covered in many large, pointed warts. [12]

In general the colour of this toad is brown or grey, mottled with irregular splotches of dark gray or black. [12] There is a thin [4] or thick, [12] lighter coloured, mid-dorsal stripe down the back. [4] [12] The dorsal colouration is light brown, often with lighter brown [4] or pale grey [12] bands toward the sides of the back. [4] [12] In females and juveniles the dorsum has darker brown mottling. The sides are dark, [4] with a broad, dark stripe below the row of lateral warts. [12] The ventral surface has a yellowish base, [4] while the throat and chest are black, [2] [12] and the underside of the lower jaw has white spots along the edge. The eyes have a tan-coloured, square-shaped patch below them, reaching to the lip. There is a mottled dark brown region from the eye to the edge of the paratoid gland, which appears somewhat like a mask. The legs are mottled with light and dark brown, [12] with dark bars on the upper surface of the thighs. [4] The juvenile is somewhat more brightly coloured. [12]

Call

The call is a "short trill" with toads having a round, inflated vocal sac. [4] The call is several seconds long, and is repeated several times, with the intervals between each trill lasting a few seconds. [12]

Similar species

In the field it is most similar to Incilius coniferus , being most easily told apart by the length of the first finger of the hand being nearly as long as the third, longest finger in the species, whereas in I. coniferus the first finger is shorter. Incilius aucoinae is very similar, but males of that species are smaller, and this species has a black chest and throat, mottling on the flanks, transverse folds between parietal crests, cranial crests that are heightened vertically, and distinct pretympanic and preorbital crests. [12]

According to Jay M. Savage (2002) the call is similar to I. luetkenii or I. valliceps. [12]

Distribution

In the 1960s the range was considered to include Nicaragua, but in 1972 Villa stated all earlier collections from this country were to be considered misidentified specimens of Incilius luetkenii or I. valliceps. [3] It was subsequently believed to be endemic to Costa Rica. [9] [6] In 2004 similar toads from the Pacific slopes of Costa Rica were reassigned to a new species, I. aucoinae. [1]

In Costa Rica it occurs throughout the northern section of the Cordillera Central at mid-latitudes, and throughout the Cordillera de Talamanca and the eastern Caribbean seaboard. As of 2008 it is unclear if I. melanochlorus also occurs on the Pacific slopes of Costa Rica, or if those are misidentified I. aucoinae. [1] McDiarmid and Savage recorded it in 2005 in meadows of the Peninsula de Osa, extending the distribution to the Pacific slopes in the far southwest of the country. [4] [12] According to Pounds et al. in 1997, it disappeared from Monteverde, Costa Rica, in the late 1980s and then reappeared there in the 1990s. [1]

In 2004 the toad was recorded in Nicaragua for first time (correctly) in the Indio Maíz Biological Reserve by Gunther Köhler and his team, [1] in 2009 it was also recorded from Río San Juan under auspices of an expedition by Köhler [4] and in 2014 it was reported in Rivas Department on the Pacific coast by a Nicaraguan team. [3] In 2009 the distribution was extended to Panama following Jaramillo and Ibáñez (2009). [3]

Ecology

This toad is nocturnal and insectivorous. [12] The toads may become more active during light rain showers. [2]

Habitat

The native habitats of this species are wet lowland rainforests and the lower premontane wet forests at elevations to about 1,080 m (3,540 ft) asl. [1] Its favourite locations are near large streams and at the edges of forests. [12] In the northeastern rainforests of Costa Rica it occurs from sea level to 2,000 m (6,600 ft) in elevation, with frogs of the genus Eleutherodactylus being the few amphibians to occur at higher altitudes here. [6] It is occasionally found amongst the leaf litter on the forest floor. [12]

Reproduction

Reproduction happens during the dry season [4] [12] starting with the males calling from January to February from pools along rocky streams, or within 50 cm (20 in) of water. Males may call at odd times of the year. Males develop dark-brown nuptial pads on their first and second fingers during breeding season. [12] Although some sources claim breeding (amplexus) takes place in large streams, [1] [4] according to Savage (2002) it occurs in small, somewhat rocky-bottomed streams which are at low water for the season. [12]

Interspecific relationships

Male toads may share their pools with the larger Bufo marinus toads. [12]

Diseases

The pathogenic fungus Batrachochytrium dendrobatidis has had no impact on this toad. [1]

Conservation

When the species had been less studied, earlier authors such as Bolaños & Chaves writing for the IUCN in 2004 mistakenly believed it to be endemic to Costa Rica. [9] [6] As such it was automatically added to the local Red List and considered threatened. It was described in 2004 as most likely not very tolerant of deforestation and siltation and pollution of its breeding habitat, but nonetheless it was downgraded to "least concern" in 2004. [9] As of 2008 the IUCN describes it as "widespread and regularly encountered" and "common and somewhat adaptable with a presumed large population". It is regularly seen during the breeding period, [1] and is common in wet forests at higher elevations at favoured habitats. [12] It occurs at La Selva Biological Station, in the Monteverde Cloud Forest Reserve (if correctly identified) [1] and throughout the Maquenque National Wildlife Refuge in Costa Rica, [6] and in the Indio Maíz Biological Reserve [1] [4] and Río San Juan Wildlife Refuge in Nicaragua. [4] It is kept in an ex situ collection at the Costa Rican Amphibian Research Center. [5]

Related Research Articles

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<span class="mw-page-title-main">Gulf Coast toad</span> Species of amphibian

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<i>Incilius aucoinae</i> Species of amphibian

Incilius aucoinae is a species of toads in the family Bufonidae. It is found in south-western Costa Rica and adjacent western Panama. Before its description in 2004, it was confused with Incilius melanochlorus.

<i>Incilius campbelli</i> Species of amphibian

Incilius campbelli is a species of toad in the family Bufonidae. It was first described in 1994. It is found in eastern Chiapas (Mexico), Guatemala, western Honduras, and Maya Mountains, Belize. Its natural habitats are lowland moist and premontane wet forests, and pristine forests in mountainous regions. It is threatened by habitat loss.

<i>Incilius coccifer</i> Species of amphibian

Incilius coccifer is a species of toad in the family Bufonidae. It is found in southern Mexico and southeastward in the Central America through Guatemala, El Salvador, Honduras, and Nicaragua to northwestern Costa Rica. Several species that were formerly included in this species have been named as distinct species: Incilius porteri, Incilius ibarrai, Incilius pisinnus, and Incilius signifer. Its natural habitats are lowland dry and moist forests, and it occurs also in disturbed areas such as pastures, roadside ditches, gardens, and vacant lots in urban areas. It is an abundant and widespread species that is not facing significant threats.

<i>Incilius coniferus</i> Species of amphibian

The evergreen toad is a species of toad in the family Bufonidae.

<i>Rhaebo haematiticus</i> Species of amphibian

Rhaebo haematiticus is a species of toad in the family Bufonidae. It is found in eastern Honduras, Nicaragua, Costa Rica, Panama, Colombia, northwestern Venezuela, and northwestern Ecuador. Its altitudinal range is from sea level to 1,300 m (4,300 ft) asl. Its natural habitats are primary tropical moist forest and submontane humid forest. It is a nocturnal, leaf-litter species that during the breeding seasons is found along small streams and large rivers. It tolerates some habitat degradation but only occurs close to forest. Threats to it are habitat loss caused by agriculture, wood extraction, and cattle ranching, and locally oil pollution and dams.

<i>Incilius ibarrai</i> Species of amphibian

Incilius ibarrai is a species of toad in the family Bufonidae. It is found in the central and southern highlands of Guatemala and adjacent Honduras. The specific name ibarrai honors Jorge Alfonso Ibarra (1921–2000), then-director of the Guatemalan National Natural History Museum.

<i>Incilius leucomyos</i> Species of amphibian

Incilius leucomyos is a species of toad in the family Bufonidae. It was described in 2000 and is endemic to the Atlantic versant of the north-central Honduras.

<i>Incilius luetkenii</i> Species of amphibian

Incilius luetkenii is a species of toad in the family Bufonidae. It is found in the Mesoamerica along the Pacific versant from central Costa Rica to extreme southern Chiapas, Mexico, as well as dry interior valleys of Guatemala and Honduras and San Juan River drainage in Costa Rica on the Atlantic versant. It occurs in open areas, including disturbed pasturelands in lowland dry forest, and to a lesser extent, in lowland moist and premontane moist forests. It breeds in temporary pools. It is a common species that is not facing major threats.

<span class="mw-page-title-main">Pine toad</span> Species of amphibian

The pine toad is a species of toad in the family Bufonidae. It is endemic to Mexico and found on the Central Mexican Plateau.

<i>Incilius pisinnus</i> Species of amphibian

Incilius pisinnus is a species of "true" toads in the family Bufonidae. It is endemic to Mexico and known from the Tepalcatepec Valley in Michoacán. Prior to its description in 2005, it was mixed with Incilius coccifer and Incilius cycladen. The specific name pisinnus, from the Latin word for "small", refers to the comparatively small size of this species among its close relatives.

Incilius epioticus is a species of toad in the family Bufonidae. It is found on the Atlantic versant of the Cordillera de Talamanca in south-eastern Costa Rica and north-eastern Panama. Its natural habitats are primary and mature secondary forests, cloud forests, and highland oak forests. It is diurnal and found over dead leaves on the forest floor.

The Latin word Cranopsis has been used to name animal genera of frogs, mollusks and branchiopods. Cranopsis, was used for an anuran, and is a junior homonym of Cranopsis, for a mollusk; and Cranopsis, for a branchiopod. Cranopsis currently describes a mollusk genus in the family Fissurellidae.

Incilius chompipe is a species of toads in the family Bufonidae, known from several localities near Cascajal in the Cerro Chompipe and in the Reserva Dantas, both in the Cordillera Central of Costa Rica.

Incilius guanacaste is a species of toads in the family Bufonidae. It is endemic to the Cordillera de Guanacaste in northern Costa Rica. The species is only known from the slopes of Miravalles Volcano and Rincón de la Vieja Volcano.

<i>Incilius</i> Genus of amphibians

Incilius is genus of toads in the true toad family, Bufonidae. They are sometimes known as the Central American toads or Middle American toads and are found in southern USA, Mexico, Central America, and northern Pacific South America. They are an ecologically and biogeographically diverse group of toads, including micro-endemic species such as Incilius spiculatus that are restricted to undisturbed cloud forests, and widespread lowland species such as Incilius valliceps that predominantly occur in disturbed habitats.

<span class="mw-page-title-main">Golden toad</span> Extinct species of toad that was endemic to Costa Rica

The golden toad is an extinct species of true toad that was once abundant in a small, high-altitude region of about 4 square kilometres (1.5 sq mi) in an area north of the city of Monteverde, Costa Rica. It was endemic to elfin cloud forest. Also called the Monte Verde toad, Alajuela toad and orange toad, it is commonly considered the "poster child" for the amphibian decline crisis. This toad was first described in 1966 by herpetologist Jay Savage. The last sighting of a single male golden toad was on 15 May 1989, and it has since been classified as extinct by the International Union for Conservation of Nature (IUCN).

References

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  2. 1 2 3 4 5 6 7 8 Taylor, Edward H. (1 July 1952). "A Review of the Frogs and Toads of Costa Rica". The University of Kansas Science Bulletin. 35 (1): 616–618. doi: 10.5962/bhl.part.4328 . Retrieved 5 September 2019.
  3. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 Frost, Darrel R. (2015). "Incilius melanochlorus (Cope, 1877)". Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History. Retrieved 3 September 2019.
  4. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 "Incilius melanochlorus Cope 1877". Amphibians of Panama. Smithsonian Tropical Research Institute. Retrieved 12 December 2015.
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  6. 1 2 3 4 5 Mauricio Salas Varga (2009). Humedales de Ramsar (FIR) – Anexo #2 Biodiversidad 2009 (PDF) (Report) (in Spanish). Centro Científico Tropical. p. 8. Retrieved 3 September 2019.
  7. Liddell, Henry George; Scott, Robert (1940). "μελανω". A Greek–English Lexicon . Oxford: Clarendon Press.
  8. Liddell, Henry George; Scott, Robert (1940). "χλωρός". A Greek–English Lexicon . Oxford: Clarendon Press.
  9. 1 2 3 4 IUCN SSC Amphibian Specialist Group (2020). "Incilius melanochlorus". IUCN Red List of Threatened Species . 2020: e.T54705A54359101. doi: 10.2305/IUCN.UK.2020-2.RLTS.T54705A54359101.en . Retrieved 17 November 2021.
  10. Frost, D. R.; Grant, T.; Faivovich, J. N.; Bain, R. H.; Haas, A.; Haddad, C. L. F. B.; De Sá, R. O.; Channing, A.; Wilkinson, M.; Donnellan, S. C.; Raxworthy, C. J.; Campbell, J. A.; Blotto, B. L.; Moler, P.; Drewes, R. C.; Nussbaum, R. A.; Lynch, J. D.; Green, D. M.; Wheeler, W. C. (2006). "The amphibian tree of life". Bulletin of the American Museum of Natural History. 297: 1–291. doi: 10.1206/0003-0090(2006)297[0001:TATOL]2.0.CO;2 . hdl:2246/5781.
  11. 1 2 Pauly, Greg B.; Hillis, David M.; Cannatella, David C. (2009). "Taxonomic freedom and the role of official lists of species names". Herpetologica. 65 (2): 115–128. doi: 10.1655/08-031R1.1 . Retrieved 4 September 2019.
  12. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 Kim, Stella; Whittaker, Kellie (2 November 2009). "Incilius melanochlorus: Wet Forest Toad". AmphibiaWeb. University of California Berkeley. Retrieved 4 September 2019.
  13. Frost, Darrel R. (2013). "Incilius Cope, 1863". Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History. Retrieved 4 July 2017.
  14. Mendelson III, Joseph R.; Mulcahy, Daniel G.; Williams, Tyler S.; Sites Jr., Jack W. (21 December 2011). "A phylogeny and evolutionary natural history of mesoamerican toads (Anura: Bufonidae: Incilius) based on morphology, life history, and molecular data". Zootaxa. 3138 (1): 1–34. doi: 10.11646/zootaxa.3138.1.1 . Retrieved 4 September 2019.