# Bolitoglossa

Bolitoglossa is a genus of lungless salamanders, also called mushroom-tongued salamanders, tropical climbing salamanders, or web-footed salamanders, in the family Plethodontidae. Their range is between northern Mexico through Central America to Colombia, Venezuela, Ecuador, Peru, northeastern Brazil, and central Bolivia. Neotropical salamanders of the Bolitoglossa make up the largest genus in the order Caudata, consisting of approximately one-fifth of all known species of salamanders. Adult salamanders range anywhere from 45mm to 200mm in length depending on their specific species. They are notorious for their ability to project their tongue at prey items, as indicated from their name. They are also known for their webbed feet, having significantly more webbing than any other species outside their genus with the exception of the cave-dwelling Mexican bolitoglossine Chiropterotriton magnipes. Although webbed feet are a common characteristic of these salamanders, only about half of the species in this genus contain webbed feet.

## Characteristics
### Hand and foot morphology

Hand and foot morphology is strikingly diverse in an otherwise morphologically uniform group. While just under half of these species contain webbing between their fingers and toes, the remaining species experience little to no webbing and undergo elongation of their fingers and toes throughout development. Ultimately, the variation of foot morphology within this genus is primarily due to natural selection. Derived characteristics correspond to arboreal vs. terrestrial salamanders.
* Webbed fingers – natural selection to improve terrestrial movement through water.
* Elongated fingers – natural selection of increased suction efficiency, favoring a larger surface area of the foot. This also selects for a decreased body size, enabling the salamander to cling trees more easily.

### Tail autotomy

Tail autotomy refers to the salamanders' ability to release or lose their tail if necessary. This is a common characteristic of nearly all salamanders and lizards. (See autotomy). It is particularly helpful to the salamander in escaping attacks from its predators. Once the tail has been lost, it can regenerate one time. After this regeneration, the tail is incapable of separation with regeneration.

### Poison

Bolitoglossa rostrata and B. subpalmata are two rare examples of poisonous salamanders within their genus. The poison is secreted through their skin as an antipredator mechanism. It is particularly toxic to certain snake species, rendering them immobile and unresponsive to external stimuli upon initial contact. The common defense tactic of these two species is to remain still in the presence of the snake until it makes initial contact (usually by the flickering of its tongue), and then run away as the paralytic poison begins to take effect in the snake.

## Evolution
### Natural selection

Tropical adaptation of the Bolitoglossa is thought to have evolved from North American plethodontids. Natural selection is responsible for morphological changes shifting from those supporting temperate environments to those supporting tropical environments such as Panama and Costa Rica. Natural selection is thought to have resulted in genetic changes from physical adaptation. The main differences that have developed from natural selection affect the skull and bones of the feet in these salamanders. Due to these primary changes, secondary changes are believed to have followed, including:
* Body size
* Additional ossification of bones
* Webbing
* Ear structure
Phylogeny of this genus is partially dependent on its variations in bone structure due to the effects of natural selection over a long period of time.

### Hybridization

The first documented case of hybridization in tropical salamanders occurred between B. frankini and B. resplendens. This hybridization has taken a pervasive effect on the morphology of B. resplendens, whereas B. frankini seemed to maintain its same physical structure.

## Taxonomy

Derived characteristics of the genus Bolitoglossa has led to their classification based on this specific list of characters:
* Tongue and hypobranchial apparatus
* Epibranchial Number
** embryos having a single epibranchial
* Tail Autotomy
* Brain stem motor control
* Bone structure of Jaws, Cranial, and inner ear
* Chromosome number
** diploid number of chromosomes is 26
* Development
Classification of this genus is primarily accomplished through analysis of the salamanders' DNA. This has proven to be the most effective and accurate way of classifying this genus.

## Species

As of January 2024, there are 140 recognised species assigned to this genus.
| Binomial Name and author | Common name |
| B. "altamazonica" Cusi, Gagliardi-Urrutia, Brcko, Wake, and von May, 2020 | |
| B. adspersa Peters, 1863 | Peter's climbing salamander |
| B. alberchi García-París, Parra-Olea, Brame & Wake, 2002 | Alberch's salamander |
| B. altamazonica Cope, 1874 | Nauta salamander |
| B. alvaradoi Taylor, 1954 | Alvarado's salamander |
| B. anthracina Brame, Savage, Wake & Hanken, 2001 | Coal-black salamander |
| B. aurae Kubicki B, Arias E. 2016 | Aura's golden salamander |
| B. aureogularis Boza-Oviedo E, Rovito SM, Chaves G, Garcia-Rodriguez A, Artavia LG, Bolanos F, Wake DB 2012 | Yellow-throated web-footed salamander |
| B. awajun Cusi JC, Gagliardi-Urrutia G, Brcko IC, Wake DB, Von May R. 2020 | Awajun Salamander |
| B. biseriata Tanner, 1962 | Two-lined climbing salamander |
| B. bolanosi Arias, Chaves, and Parra-Olea, 2024 | Bolaños’ Web-footed Salamander |
| B. borburata Trapido, 1942 | Carabobo climbing salamander |
| B. bramei Wake, Savage & Hanken, 2007 | Brame's climbing salamander |
| B. caldwellae Brcko IC, Hoogmoed MS, Neckel-Oliveira S 2013 | Caldwell's mushroomtongue salamander |
| B. capitana Brame & Wake, 1963 | Orphan salamander |
| B. carri McCranie & Wilson, 1993 | Cloud forest salamander |
| B. cathyledecae Ponce M, Navarro D, Morales R, and Batista A., 2022 | Chiriquí fire salamander |
| B. cataguana Townsend, Butler, Wilson & Austin, 2009 | Cataguana salamander |
| B. celaque McCranie & Wilson, 1993 | Celaque climbing salamander |
| B. centenorum Campbell et al., 2010 | |
| B. cerroensis Taylor, 1952 | Millville climbing salamander |
| B. chica Brame & Wake, 1963 | Hotel Zaracay salamander |
| B. chinanteca Rovito, Parra-Olea, Lee & Wake, 2012 | Chinanteca salamander |
| B. chucantiensis Batista A, Koehler G, Mebert K, Vesely M 2014 | Chucanti salamander |
| B. coaxtlahuacana Palacios-Aguilar R, Cisneros-Bernal AY, Arias-Montiel JD, Parra-Olea G. 2020 | Coaxtlahuacán Salamander |
| B. colonnea Dunn, 1924 | La Loma salamander |
| B. compacta Wake, Brame & Duellman, 1973 | Cerro Pando salamander |
| B. conanti McCranie & Wilson, 1993 | Conant's salamander |
| B. copia Wake, Hanken & Ibáñez, 2005 Archived 12 March 2012 at the Wayback Machine | El Cope giant salamander |
| B. copinhorum Itgen MW, Sessions SK, Wilson LD, Townsend JH 2020 | Lenca mushroomtongue salamander |
| B. cuchumatana Stuart, 1943 | Oak forest salamander |
| B. cuna Wake, Brame, & Duellman, 1973 | Camp Sasardi salamander |
| B. daryorum Campbell et al., 2010 | |
| B. decora McCranie & Wilson, 1997 | Monte Escondido salamander |
| B. diaphora McCranie & Wilson, 1995 | El Cusuco salamander |
| B. digitigrada Wake, Brame & Thomas, 1982 | Rio Santa Rosa salamander |
| B. diminuta Robinson, 1976 | Quebrada Valverde salamander |
| B. dofleini Werner, 1903 | Doflein's salamander |
| B. dunni Schmidt, 1933 | Dunn's climbing salamander |
| B. engelhardti Schmidt, 1936 | Engelhardt's climbing salamander |
| B. epimela Wake & Brame, 1963 | Tapanti climbing salamander |
| B. equatoriana Brame & Wake, 1972 | Ecuadorian climbing salamander |
| B. eremia Campbell et al., 2010 | |
| B. flavimembris Schmidt, 1936 | Yellow-legged climbing salamander |
| B. flaviventris Schmidt, 1936 | Yellow-belly climbing salamander |
| B. franklini Schmidt, 1936 | Franklin's climbing salamander |
| B. gomezi Wake, Savage & Hanken, 2007 | Gómez's web-footed salamander |
| B. gracilis Bolaños, Robinson & Wake, 1987 | Rio Quiri salamander |
| B. guaneae Acosta-Galvis AR, Gutierrez-Lamus DL 2012 | |
| B. guaramacalensis Schargel, García-Pérez & Smith, 2002 | Guaramacal salamander |
| B. hartwegi Wake & Brame, 1969 | Hartweg's climbing Salamander |
| B. heiroreias Greenbaum, 2004 | Holy-Mountain salamander |
| B. helmrichi Schmidt, 1836 | Coban climbing salamander |
| B. hermosa Papenfuss, Wake & Adler, 1984 | Guerreran climbing salamander |
| B. hiemalis Lynch, 2001 | Winter climbing salamander |
| B. huehuetenanguensis Campbell et al., 2010 | |
| B. hypacra Brame & Wake, 1962 | Paramo Frontino salamander |
| B. indio Wake, et al., 2008 | Rio Indio Webfoot Salamander |
| B. insularis Wake, et al., 2008 | |
| B. jacksoni Elias, 1984 | Jackson's climbing salamander |
| B. jugivagans Hertz A, Lotzkat S, Koehler G 2013 | Continental divide salamander |
| B. kamuk Boza-Oviedo, Rovito, Chaves, García-Rodríguez, Artavia, Bolaños, and Wake, 2012 | Kamuk Web-footed Salamander |
| B. kaqchikelorum Campbell et al., 2010 | |
| B. la Campbell et al., 2010 | |
| B. leandrae Acevedo AA, Wake DB, Marquez R, Silva K, Franco R, Amezquita A 2013 | Leandra salamander |
| B. lignicolor Peters, 1873 | Camron climbing salamander |
| B. lincolni Stuart, 1943 | Lincoln's climbing salamander |
| B. longissima McCranie & Cruz, 1996 | Longest climbing salamander |
| B. lozanoi Acosta-Galvis & Restrepo, 2001 | Lozano's salamander |
| B. macrinii Lafrentz, 1930 | Oaxacan climbing salamander |
| Binomial Name and Author | Common Name |
| B. madeira Brcko IC, Hoogmoed MS, Neckel-Oliveira S 2013 | |
| B. magnifica Hanken, Wake & Savage, 2005 | Magnificent web-footed salamander |
| B. marmorea Tanner & Brame, 1961 | Crater salamander |
| B. medemi Brame & Wake, 1972 | Finca Chibigui salamander |
| B. meliana Wake & Lynch, 1982 | Meliana climbing salamander |
| B. mexicana Duméril, Bibron & Duméril, 1854 | Mexican climbing salamander |
| B. minutula Wake, Brame & Duellman, 1973 | Dwarf climbing salamander |
| B. mombachoensis Köhler & McCranie, 1999 | Mombacho salamander |
| B. morio Cope, 1869 | Cope's climbing salamander |
| B. mucuyensis Garcia-Gutierrez J, Escalona M, Mora A, Diaz de Pascual A, Fermin G. 2013 | La Mucuy salamander |
| B. muisca López-Perilla, Fernández-Roldán, Meza-Joya, and Medina-Rangel, 2023 | Muisca salamander |
| B. mulleri Brocchi, 1883 | Muller's climbing salamander |
| B. nicefori Brame & Wake, 1963 | San Gil climbing salamander |
| B. nigrescens Taylor, 1949 | Cordillera central salamander |
| B. ninadormida Campbell et al., 2010 | |
| B. nussbaumi Campbell et al., 2010 | |
| B. nympha Campbell et al., 2010 | |
| B. oaxacensis Parra-Olea, Garcia-Paris & Wake, 2002 | Atoyac web-footed salamander |
| B. obscura Hanken, Wake & Savage, 2005 | Tapantí giant salamander |
| B. occidentalis Taylor, 1941 | Southern banana salamander |
| B. odonnelli Stuart, 1943 | O'Donnell's climbing salamander |
| B. omniumsanctorum Stuart, 1952 | Todos Santos Salamander |
| B. oresbia McCranie, Espinal & Wilson, 2005 Archived 16 March 2012 at the Wayback Machine | Zarciadero web-footed salamander |
| B. orestes Brame & Wake, 1962 | Culata climbing salamander |
| B. pacaya Campbell et al., 2010 | |
| B. palmata Werner, 1897 | Amazon climbing salamander |
| B. pandi Brame & Wake, 1963 | Pandi climbing salamander |
| B. paraensis Unterstein, 1930 | Para climbing salamander |
| B. peruviana Boulenger, 1883 | Peruvian climbing salamander |
| B. pesrubra Taylor, 1952 | Red-footed climbing salamander |
| B. phalarosoma Wake & Brame, 1962 | Medellin climbing salamander |
| B. platydactyla Gray in Cuvier, 1831 | Broadfoot climbing salamander |
| B. porrasorum McCranie & Wilson, 1995 | Pijol salamander |
| B. psephena Campbell et al., 2010 | |
| B. pygmaea Bolaños & Wake, 2009 | Pygmy web-footed salamander |
| B. qeqom Hellen D-B, Serrano MJ, Alonso-Ascencio M, Cruz-Font JJ, Rosito-Prado I, Ruiz-Villanueva KJA, Vásquez-Almazán JR, Ariano-Sánchez D. 2021 | |
| B. ramosi Brame & Wake, 1972 | Ramos' climbing salamander |
| B. riletti Holman, 1964 | Rilett's climbing salamander |
| B. robinsoni Bolaños & Wake, 2009 | Robinson's web-footed salamander |
| B. robusta Cope, 1894 | Robust climbing salamander |
| B. rostrata Brocchi, 1883 | Long-nosed climbing salamander |
| B. rufescens Cope, 1869 | Northern banana Salamander |
| B. salvinii Gray, 1868 | Salvin's salamander |
| B. savagei Brame & Wake, 1963 | Savage's salamander |
| B. schizodactyla Wake & Brame, 1966 | Cocle salamander |
| B. silverstonei Brame & Wake, 1972 | Silverstone's salamander |
| B. sima Vaillant, 1911 | Northwestern climbing salamander |
| B. sombra Hanken, Wake & Savage, 2005 | Shadowy web-footed salamander |
| B. sooyorum Vial, 1963 | Cordillera Talamanca salamander |
| B. spendida Boza-Oviedo, Rovito, Chaves, García-Rodríguez, Artavia, Bolaños, and Wake, 2012 | Splendid Web-footed Salamander |
| B. striatula Noble, 1918 | Cukra climbing salamander |
| B. stuarti Wake & Brame, 1969 | Stuart's salamander |
| B. subpalmata Boulenger, 1896 | La Palma salamander |
| B. suchitanensis Campbell et al., 2010 | |
| B. synoria McCranie & Köhler, 1999 | Cerro Pital salamander |
| B. tamaense Acevedo AA, Wake DB, Marquez R, Silva K, Franco R, Amezquita A 2013 | Tama salamander |
| B. tapajonica Brcko IC, Hoogmoed MS, Neckel-Oliveira S 2013 | |
| B. tatamae Acosta-Galvis & Hoyos, 2006 | Tatama climbing salamander |
| B. taylori Wake, Brame & Myers, 1970 | Taylor's salamander |
| B. tenebrosa Vasquez-Almazan CR, Rovito SM 2014 | Guatemalan black salamander |
| B. tica García-París, Parra-Olea & Wake, 2008 | Tico salamander |
| B. tzultacaj Campbell et al., 2010 | |
| B. vallecula Brame & Wake, 1963 | Yarumal climbing salamander |
| B. veracrucis Taylor, 1951 | Veracruz salamander |
| B. walkeri Brame & Wake, 1972 | Walker's salamander |
| B. xibalba Campbell et al., 2010 | |
| B. yariguiensis Meza-Joya FL, Hernandez-Jaimes C, Ramos-Pallares E. 2017 | Yariguíes salamander |
| B. yucatana Peters, 1882 | Yucatán salamander |
| B. zacapensis Rovito, Vásquez-Almazán & Papenfuss, 2010 | Zacapa salamander |
| B. zapoteca Parra-Olea, Garcia-Paris & Wake, 2002 | Zapotec salamander |