Glass frog
Encyclopedia
Glass frog is the common name for the frog
Frog
Frogs are amphibians in the order Anura , formerly referred to as Salientia . Most frogs are characterized by a short body, webbed digits , protruding eyes and the absence of a tail...

s of the amphibian
Amphibian
Amphibians , are a class of vertebrate animals including animals such as toads, frogs, caecilians, and salamanders. They are characterized as non-amniote ectothermic tetrapods...

 family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...

 Centrolenidae (order
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...

 Anura). While the general background coloration of most glass frogs is primarily lime green
Lime (color)
Lime, also traditionally known as lime green or lime-green, is a color three-quarters of the way between yellow and green , so named because it is a representation of the color of the citrus fruit called limes...

, the abdominal skin of some members of this family is transparent
Transparency and translucency
In the field of optics, transparency is the physical property of allowing light to pass through a material; translucency only allows light to pass through diffusely. The opposite property is opacity...

. The internal viscera, including the heart
Heart
The heart is a myogenic muscular organ found in all animals with a circulatory system , that is responsible for pumping blood throughout the blood vessels by repeated, rhythmic contractions...

, liver
Liver
The liver is a vital organ present in vertebrates and some other animals. It has a wide range of functions, including detoxification, protein synthesis, and production of biochemicals necessary for digestion...

, and gastrointestinal tract
Gastrointestinal tract
The human gastrointestinal tract refers to the stomach and intestine, and sometimes to all the structures from the mouth to the anus. ....

 are visible through this translucent skin, hence the common name.

Taxonomy

The first described species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

 of Centrolenidae was the "giant" Centrolene
Centrolene
Centrolene is a genus of glass frogs in the family Centrolenidae. The adult males are characterized by having a humeral spine, as most members of this family...

 geckoideum
, named by Marcos Jiménez de la Espada
Marcos Jiménez de la Espada
Marcos Jiménez de la Espada was a Spanish zoologist, explorer and writer, born in Cartagena, Spain, although he spent most of his life in Madrid, where he died. He is known for participating in the Pacific Scientific Commission, with whom he traveled America from 1862 to 1865...

 in 1872, based on a specimen collected in northeastern Ecuador
Ecuador
Ecuador , officially the Republic of Ecuador is a representative democratic republic in South America, bordered by Colombia on the north, Peru on the east and south, and by the Pacific Ocean to the west. It is one of only two countries in South America, along with Chile, that do not have a border...

. Several species were described in subsequent years by different herpetologists (including G. A. Boulenger
George Albert Boulenger
George Albert Boulenger FRS was a Belgian-British zoologist who identified over 2000 new animal species, chiefly fish, reptiles and amphibians.-Life:...

, G. K. Noble, and E. H. Taylor
Edward Harrison Taylor
Edward Harrison Taylor was an American herpetologist from Kansas.He was born in Maysville, Missouri and studied at the University of Kansas in Lawrence, Kansas, graduating with a B.A. in 1912. Subsequently, he went to the Philippines, where at first he held a teacher's post in a village in central...

) but usually placed together with the tree frog
Tree frog
Hylidae is a wide-ranging family of frogs commonly referred to as "tree frogs and their allies". However, the hylids include a diversity of frog species, many of which do not live in trees, but are terrestrial or semi-aquatic.-Characteristics:...

s in the genera Hylella or Hyla
Hyla
The genus Hyla is one of approximately 38 genera in the New World family of tree frogs . The word Hyla translates to "tree," and tree frogs are indeed arboreal...

.

The family Centrolenidae was proposed by Edward H. Taylor
Edward Harrison Taylor
Edward Harrison Taylor was an American herpetologist from Kansas.He was born in Maysville, Missouri and studied at the University of Kansas in Lawrence, Kansas, graduating with a B.A. in 1912. Subsequently, he went to the Philippines, where at first he held a teacher's post in a village in central...

 in 1951. Between the '50s and '70s, most species of glass frogs were known from Central America
Central America
Central America is the central geographic region of the Americas. It is the southernmost, isthmian portion of the North American continent, which connects with South America on the southeast. When considered part of the unified continental model, it is considered a subcontinent...

, particularly from Costa Rica
Costa Rica
Costa Rica , officially the Republic of Costa Rica is a multilingual, multiethnic and multicultural country in Central America, bordered by Nicaragua to the north, Panama to the southeast, the Pacific Ocean to the west and the Caribbean Sea to the east....

 and Panama
Panama
Panama , officially the Republic of Panama , is the southernmost country of Central America. Situated on the isthmus connecting North and South America, it is bordered by Costa Rica to the northwest, Colombia to the southeast, the Caribbean Sea to the north and the Pacific Ocean to the south. The...

, where E. H. Taylor and Jay M. Savage extensively worked, and just a few species were known to occur in South America
South America
South America is a continent situated in the Western Hemisphere, mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere. The continent is also considered a subcontinent of the Americas. It is bordered on the west by the Pacific Ocean and on the north and east...

. In 1973, John D. Lynch and William E. Duellman, published a large revision of the glass frogs from Ecuador showing that the species richness of Centrolenidae was particularly concentrated in the Andes
Andes
The Andes is the world's longest continental mountain range. It is a continual range of highlands along the western coast of South America. This range is about long, about to wide , and of an average height of about .Along its length, the Andes is split into several ranges, which are separated...

. Later contributions by authors like Juan Rivero, Jay Savage, William Duellman, John D. Lynch, Pedro Ruiz-Carranza and José Ayarzagüena increased the number of described taxa especially from Central America, Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

, Colombia
Colombia
Colombia, officially the Republic of Colombia , is a unitary constitutional republic comprising thirty-two departments. The country is located in northwestern South America, bordered to the east by Venezuela and Brazil; to the south by Ecuador and Peru; to the north by the Caribbean Sea; to the...

, Ecuador
Ecuador
Ecuador , officially the Republic of Ecuador is a representative democratic republic in South America, bordered by Colombia on the north, Peru on the east and south, and by the Pacific Ocean to the west. It is one of only two countries in South America, along with Chile, that do not have a border...

, and Peru
Peru
Peru , officially the Republic of Peru , is a country in western South America. It is bordered on the north by Ecuador and Colombia, on the east by Brazil, on the southeast by Bolivia, on the south by Chile, and on the west by the Pacific Ocean....

.

The evolutionary relationships, biogeography, and character evolution of centrolenids were discussed by Guayasamin et al. (2008). Glass frogs originated in South America and dispersed multiple times into Central America. Character evolution seems to be complex, with multiple gains and/or losses of humeral spines, reduced hand webbing, and complete ventral transparency.

The taxonomical classification of the glass frogs has been problematic. In 1991, after a major revision of the species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

 and taxonomic characters, the herpetologists Pedro Ruiz-Carranza and John D. Lynch published a proposal for a taxonomic classification of the Centrolenidae based on cladistic
Cladistics
Cladistics is a method of classifying species of organisms into groups called clades, which consist of an ancestor organism and all its descendants . For example, birds, dinosaurs, crocodiles, and all descendants of their most recent common ancestor form a clade...

 principles and defining monophyletic
Monophyly
In common cladistic usage, a monophyletic group is a taxon which forms a clade, meaning that it contains all the descendants of the possibly hypothetical closest common ancestor of the members of the group. The term is synonymous with the uncommon term holophyly...

 groups. That paper was the first of a series of contributions dealing with the glass frogs from Colombia that lead them to described almost 50 species of glass frogs. The genus Centrolene was proposed to include the species with a humeral spine
Humeral spine
Humeral spines are ventrolateral extensions of the humeral crista ventralis. This structures are present in the humerus of some frogs . The majority of anuran species that present humeral spines are glassfrogs but humeral spines have been reported in various other species of different families of...

 in adult males, and the genus Hyalinobatrachium
Hyalinobatrachium
Hyalinobatrachium is a genus of glassfrogs characterized by having a bulbous liver covered by white pigment, a transparent parietal peritoneum, extensive membrane between the outer fingers, and lacking a humeral spine in adult males....

to include the species with a bulbous liver. However, there was an heterogeneous group of species that they left in the genus Cochranella
Cochranella
Cochranella is a genus of glass frogs, characterized by lacking humeral spines in males, and having a lobed liver. About one-third of the species formerly placed in Cochranella were placed into the genus Nymphargus....

, defined just by lacking a humeral spine
Humeral spine
Humeral spines are ventrolateral extensions of the humeral crista ventralis. This structures are present in the humerus of some frogs . The majority of anuran species that present humeral spines are glassfrogs but humeral spines have been reported in various other species of different families of...

 and a bulbous liver. Since the publication of the extensive revision of the Colombian glass frogs, several other publications have dealt with the glass frogs from Venezuela, Costa Rica, and Ecuador.

In 2006, the genus Nymphargus
Nymphargus
Nymphargus is a newly established genus of glass frogs, characterized by lacking webbing among the outer fingers, lacking humeral spines in adult males, and having a lobed liver covered by a transparent hepatic peritoneum.-Species:...

was erected for the species with basal webbing among outer fingers (part of the previous Cochranella ocellata species group).

The four genera (Centrolene, Cochranella, Hyalinobatrachium, Nymphargus) have been shown to be poly-
Polyphyly
A polyphyletic group is one whose members' last common ancestor is not a member of the group.For example, the group consisting of warm-blooded animals is polyphyletic, because it contains both mammals and birds, but the most recent common ancestor of mammals and birds was cold-blooded...

 or paraphyletic
Paraphyly
A group of taxa is said to be paraphyletic if the group consists of all the descendants of a hypothetical closest common ancestor minus one or more monophyletic groups of descendants...

 (2008) and recently a new taxonomy has been proposed (see below).

Classification

The family Centrolenidae is a clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

 of anurans
Frog
Frogs are amphibians in the order Anura , formerly referred to as Salientia . Most frogs are characterized by a short body, webbed digits , protruding eyes and the absence of a tail...

. Previously, the family Centrolenidae was considered closely related to the family Hylidae; however, recent phylogenetic
Phylogenetics
In biology, phylogenetics is the study of evolutionary relatedness among groups of organisms , which is discovered through molecular sequencing data and morphological data matrices...

 studies have placed the Centrolenidae (and its sister taxon
Taxon
|thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement...

, the family Allophrynidae, closer to the family Leptodactylidae
Leptodactylidae
Leptodactylidae is a diverse family of frogs that probably diverged from other hyloids during the Cenozoic era, or possibly at the end of the Mesozoic. There are roughly 50 genera, one of which is Eleutherodactylus, the largest vertebrate genus, with over 700 species...

.

The monophyly
Monophyly
In common cladistic usage, a monophyletic group is a taxon which forms a clade, meaning that it contains all the descendants of the possibly hypothetical closest common ancestor of the members of the group. The term is synonymous with the uncommon term holophyly...

 of Centrolenidae is supported by morphological and behavioral characters including: (1) presence of a dilated process on the medial side of the third metacarpal (an apparently unique synapomorphy
Synapomorphy
In cladistics, a synapomorphy or synapomorphic character is a trait that is shared by two or more taxa and their most recent common ancestor, whose ancestor in turn does not possess the trait. A synapomorphy is thus an apomorphy visible in multiple taxa, where the trait in question originates in...

); (2) ventral origin of the musculus flexor teres digiti III relative to the musculus transversi metacarpi I; (3) terminal phalanges T-shaped; (4) exotroph, lotic, burrower/fossorial tadpoles with a vermiform body and dorsal C-shaped eyes, that live buried within leaf packs in still or flowing water systems; (5) eggs clutches deposited outside of water on vegetation or rocks above still or flowing water systems. Several molecular synapomorphies
Synapomorphy
In cladistics, a synapomorphy or synapomorphic character is a trait that is shared by two or more taxa and their most recent common ancestor, whose ancestor in turn does not possess the trait. A synapomorphy is thus an apomorphy visible in multiple taxa, where the trait in question originates in...

 also support the monophyly of the clade.

The taxonomic classification of Centrolenidae was recently modified. The family Centrolenidae now contains two subfamilies and twelve genera (Guayasamin et al., 2009).

Genera

Family CENTROLENIDAE
  • Subfamily Centroleninae
    • Genus Centrolene
      Centrolene
      Centrolene is a genus of glass frogs in the family Centrolenidae. The adult males are characterized by having a humeral spine, as most members of this family...

      Jiménez de la Espada, 1872
    • Genus Chimerella Guayasamin, Castroviejo, Trueb, Ayarzagüena, Rada, Vilá, 2009
    • Genus Cochranella
      Cochranella
      Cochranella is a genus of glass frogs, characterized by lacking humeral spines in males, and having a lobed liver. About one-third of the species formerly placed in Cochranella were placed into the genus Nymphargus....

      Taylor, 1951
    • Genus Espadarana Guayasamin, Castroviejo, Trueb, Ayarzagüena, Rada, Vilá, 2009
    • Genus Nymphargus
      Nymphargus
      Nymphargus is a newly established genus of glass frogs, characterized by lacking webbing among the outer fingers, lacking humeral spines in adult males, and having a lobed liver covered by a transparent hepatic peritoneum.-Species:...

      Cisneros-Heredia & McDiarmid, 2007
    • Genus Rulyrana Guayasamin, Castroviejo, Trueb, Ayarzagüena, Rada, Vilá, 2009
    • Genus Sachatamia Guayasamin, Castroviejo, Trueb, Ayarzagüena, Rada, Vilá, 2009
    • Genus Teratohyla Taylor, 1951
    • Genus Vitreorana Guayasamin, Castroviejo, Trueb, Ayarzagüena, Rada, Vilá, 2009
  • Subfamily Hyalinobatrachinae
    • Genus Celsiella Guayasamin, Castroviejo, Trueb, Ayarzagüena, Rada, Vilá, 2009
    • Genus Hyalinobatrachium
      Hyalinobatrachium
      Hyalinobatrachium is a genus of glassfrogs characterized by having a bulbous liver covered by white pigment, a transparent parietal peritoneum, extensive membrane between the outer fingers, and lacking a humeral spine in adult males....

      Ruiz-Carranza & Lynch, 1991 - "True" Glass Frogs
  • Subfamily Allophryninae

Incertae sedis Ikakogi Guayasamin, Castroviejo, Trueb, Ayarzagüena, Rada, Vilá, 2009

Characteristics

Glass frogs are generally small, ranging from 3 to 7.5 cm (1.2 to 3 in) in length. The frog is known to eat its own young. They are green in color over most of their bodies, save for the skin along the lower surface of the body, which is translucent.

Glass frogs are similar in appearance to some green frogs of the genus Eleutherodactylus
Eleutherodactylus
Eleutherodactylus is a genus of frogs in the Leptodactylidae family. It is typically described as the largest vertebrate genus on Earth, with over 700 species...

and to some tree frog
Tree frog
Hylidae is a wide-ranging family of frogs commonly referred to as "tree frogs and their allies". However, the hylids include a diversity of frog species, many of which do not live in trees, but are terrestrial or semi-aquatic.-Characteristics:...

s of the family Hylidae. However, hylid tree frogs have eyes that face to the side, whilst those of glass frogs face forward. Some species of green tree frogs (especially juveniles), such as Hyloscirtus
Hyloscirtus
Hyloscirtus is a genus of frogs in the Hylidae family. This genus was resurrected in 2005 following a major revision of the Hylidae family. The distinguishing feature being 56 transformations in nuclear and mitochondrial protein and ribosomal genes. 28 species that were previously placed in the...

 palmeri
and Hypsiboas
Hypsiboas
Hypsiboas is a genus of frogs in the Hylidae family. This genus was resurrected in 2005 following a major revision of the Hylidae family. 70 species that were previously placed in the Hyla genus were moved to this genus.-Species:...

 pellucens
, have the transparent abdominal skin typical of glass frogs, but they also have calcar
Calcar
The calcar, also known as the calcaneum , is the name given to a spur of cartilage arising from inner side of ankle and running along part of outer interfemoral membrane in bats , this is to help spread the interfemoral membrane , which is part of the wing membrane between the tail and the hind...

s on the heels, a character not present in any species of the family Centrolenidae.

Distribution

Centrolenidae is a diverse type of frogs distributed from southern Mexico
Mexico
The United Mexican States , commonly known as Mexico , is a federal constitutional republic in North America. It is bordered on the north by the United States; on the south and west by the Pacific Ocean; on the southeast by Guatemala, Belize, and the Caribbean Sea; and on the east by the Gulf of...

 to Panama
Panama
Panama , officially the Republic of Panama , is the southernmost country of Central America. Situated on the isthmus connecting North and South America, it is bordered by Costa Rica to the northwest, Colombia to the southeast, the Caribbean Sea to the north and the Pacific Ocean to the south. The...

, and through the Andes
Andes
The Andes is the world's longest continental mountain range. It is a continual range of highlands along the western coast of South America. This range is about long, about to wide , and of an average height of about .Along its length, the Andes is split into several ranges, which are separated...

 from Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...

 and the island of Tobago
Tobago
Tobago is the smaller of the two main islands that make up the Republic of Trinidad and Tobago. It is located in the southern Caribbean, northeast of the island of Trinidad and southeast of Grenada. The island lies outside the hurricane belt...

 to Bolivia
Bolivia
Bolivia officially known as Plurinational State of Bolivia , is a landlocked country in central South America. It is the poorest country in South America...

, with some species in the Amazon
Amazon River
The Amazon of South America is the second longest river in the world and by far the largest by waterflow with an average discharge greater than the next seven largest rivers combined...

 and Orinoco River basins, the Guiana Shield region, southeastern Brazil
Brazil
Brazil , officially the Federative Republic of Brazil , is the largest country in South America. It is the world's fifth largest country, both by geographical area and by population with over 192 million people...

, and northern Argentina
Argentina
Argentina , officially the Argentine Republic , is the second largest country in South America by land area, after Brazil. It is constituted as a federation of 23 provinces and an autonomous city, Buenos Aires...

.

Biology

Glass frogs are mostly arboreal. They live along rivers and streams during the breeding season, and are particularly diverse in montane cloud forest
Cloud forest
A cloud forest, also called a fog forest, is a generally tropical or subtropical evergreen montane moist forest characterized by a persistent, frequent or seasonal low-level cloud cover, usually at the canopy level. Cloud forests often exhibit an abundance of mosses covering the ground and...

s of Central and South America, although some species occur also in Amazon
Amazon Rainforest
The Amazon Rainforest , also known in English as Amazonia or the Amazon Jungle, is a moist broadleaf forest that covers most of the Amazon Basin of South America...

 and Chocóan
Tumbes-Chocó-Magdalena
Tumbes-Chocó-Magdalena is a biodiversity hotspot, which includes the tropical moist forests and tropical dry forests of the Pacific coast of South America and the Galapagos Islands. The region extends from easternmost Panama to the lower Magdalena Valley of Colombia, and along the Pacific coast of...

 rainforest
Rainforest
Rainforests are forests characterized by high rainfall, with definitions based on a minimum normal annual rainfall of 1750-2000 mm...

 and semi-deciduous
Semi-deciduous
Semi-deciduous is a botanical term which refers to plants that lose part of their foliage, also Semi-deciduous plants can mean, that plants can lose their foliage for a very short period, when old leaves fall off and new foliage growth is starting. This phenomenon occurs in tropical and...

 forests.

The eggs are usually deposited on the leaves of trees or shrubs hanging over the running water of mountain streams, creeks, and small rivers. One species leave its eggs over stones close to waterfalls. The method of egg-laying on the leaf varies between species. The males usually call from leaves close to their egg clutches. The eggs are less vulnerable to predators than those laid within water, but are affected by the parasitic maggot
Maggot
In everyday speech the word maggot means the larva of a fly ; it is applied in particular to the larvae of Brachyceran flies, such as houseflies, cheese flies, and blowflies, rather than larvae of the Nematocera, such as mosquitoes and Crane flies...

s of some fly species. As a result, some glass frogs show parental care. After they hatch, the tadpole
Tadpole
A tadpole or polliwog is the wholly aquatic larval stage in the life cycle of an amphibian, particularly that of a frog or toad.- Appellation :...

s fall into the waters below. The tadpoles are elongated, with powerful tails and low fins, suited for fast flowing water. Outside of the breeding season some species live in the canopy
Canopy (forest)
In biology, the canopy is the aboveground portion of a plant community or crop, formed by plant crowns.For forests, canopy also refers to the upper layer or habitat zone, formed by mature tree crowns and including other biological organisms .Sometimes the term canopy is used to refer to the extent...

.

External links

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