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The western clawed frog (''Xenopus tropicalis'') is a species of frog in the family
Pipidae The Pipidae are a family of primitive, tongueless frogs. The 41 species in the family Pipidae are found in tropical South America (genus ''Pipa'') and sub-Saharan Africa (the three other genera). Description Pipid frogs are highly aquatic and ...
, also known as tropical clawed frog. It is the only species in the genus ''
Xenopus ''Xenopus'' () (Gk., ξενος, ''xenos''=strange, πους, ''pous''=foot, commonly known as the clawed frog) is a genus of highly aquatic frogs native to sub-Saharan Africa. Twenty species are currently described within it. The two best-know ...
'' to have a
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectiv ...
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
. Its genome has been sequenced, making it a significant
model organism A model organism (often shortened to model) is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workin ...
for
genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar work ...
that complements the related species ''Xenopus laevis'' (the
African clawed frog The African clawed frog (''Xenopus laevis'', also known as the xenopus, African clawed toad, African claw-toed frog or the ''platanna'') is a species of African aquatic frog of the family Pipidae. Its name is derived from the three short claws o ...
), a widely used
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
model for
developmental biology Developmental biology is the study of the process by which animals and plants grow and develop. Developmental biology also encompasses the biology of regeneration, asexual reproduction, metamorphosis, and the growth and differentiation of st ...
. ''X. tropicalis'' also has a number of advantages over ''X. laevis'' in research, such as a much shorter
generation time In population biology and demography, generation time is the average time between two consecutive generations in the lineages of a population. In human populations, generation time typically ranges from 22 to 33 years. Historians sometimes use this ...
(<5 months), smaller size ( body length), and a larger number of eggs per
spawn Spawn or spawning may refer to: * Spawn (biology), the eggs and sperm of aquatic animals Arts, entertainment, and media * Spawn (character), a fictional character in the comic series of the same name and in the associated franchise ** '' Spawn: A ...
. It is found in
Benin Benin ( , ; french: Bénin , ff, Benen), officially the Republic of Benin (french: République du Bénin), and formerly Dahomey, is a country in West Africa. It is bordered by Togo to the west, Nigeria to the east, Burkina Faso to the nort ...
,
Burkina Faso Burkina Faso (, ; , ff, 𞤄𞤵𞤪𞤳𞤭𞤲𞤢 𞤊𞤢𞤧𞤮, italic=no) is a landlocked country in West Africa with an area of , bordered by Mali to the northwest, Niger to the northeast, Benin to the southeast, Togo and Ghana t ...
,
Cameroon Cameroon (; french: Cameroun, ff, Kamerun), officially the Republic of Cameroon (french: République du Cameroun, links=no), is a country in west-central Africa. It is bordered by Nigeria to the west and north; Chad to the northeast; the ...
,
Ivory Coast Ivory Coast, also known as Côte d'Ivoire, officially the Republic of Côte d'Ivoire, is a country on the southern coast of West Africa. Its capital is Yamoussoukro, in the centre of the country, while its largest city and economic centre i ...
,
Equatorial Guinea Equatorial Guinea ( es, Guinea Ecuatorial; french: Guinée équatoriale; pt, Guiné Equatorial), officially the Republic of Equatorial Guinea ( es, link=no, República de Guinea Ecuatorial, french: link=no, République de Guinée équatoria ...
,
Gambia The Gambia,, ff, Gammbi, ar, غامبيا officially the Republic of The Gambia, is a country in West Africa. It is the smallest country within mainland AfricaHoare, Ben. (2002) ''The Kingfisher A-Z Encyclopedia'', Kingfisher Publicatio ...
,
Ghana Ghana (; tw, Gaana, ee, Gana), officially the Republic of Ghana, is a country in West Africa. It abuts the Gulf of Guinea and the Atlantic Ocean to the south, sharing borders with Ivory Coast in the west, Burkina Faso in the north, and Tog ...
,
Guinea Guinea ( ),, fuf, 𞤘𞤭𞤲𞤫, italic=no, Gine, wo, Gine, nqo, ߖߌ߬ߣߍ߫, bm, Gine officially the Republic of Guinea (french: République de Guinée), is a coastal country in West Africa. It borders the Atlantic Ocean to the we ...
,
Guinea-Bissau Guinea-Bissau ( ; pt, Guiné-Bissau; ff, italic=no, 𞤘𞤭𞤲𞤫 𞤄𞤭𞤧𞤢𞥄𞤱𞤮, Gine-Bisaawo, script=Adlm; Mandinka: ''Gine-Bisawo''), officially the Republic of Guinea-Bissau ( pt, República da Guiné-Bissau, links=no ) ...
,
Liberia Liberia (), officially the Republic of Liberia, is a country on the West African coast. It is bordered by Sierra Leone to Liberia–Sierra Leone border, its northwest, Guinea to Guinea–Liberia border, its north, Ivory Coast to Ivory Coast� ...
,
Nigeria Nigeria ( ), , ig, Naìjíríyà, yo, Nàìjíríà, pcm, Naijá , ff, Naajeeriya, kcg, Naijeriya officially the Federal Republic of Nigeria, is a country in West Africa. It is situated between the Sahel to the north and the Gulf o ...
,
Senegal Senegal,; Wolof: ''Senegaal''; Pulaar: 𞤅𞤫𞤲𞤫𞤺𞤢𞥄𞤤𞤭 (Senegaali); Arabic: السنغال ''As-Sinighal'') officially the Republic of Senegal,; Wolof: ''Réewum Senegaal''; Pulaar : 𞤈𞤫𞤲𞤣𞤢𞥄𞤲𞤣𞤭 ...
,
Sierra Leone Sierra Leone,)]. officially the Republic of Sierra Leone, is a country on the southwest coast of West Africa. It is bordered by Liberia to the southeast and Guinea surrounds the northern half of the nation. Covering a total area of , Sierr ...
,
Togo Togo (), officially the Togolese Republic (french: République togolaise), is a country in West Africa. It is bordered by Ghana to the west, Benin to the east and Burkina Faso to the north. It extends south to the Gulf of Guinea, where its c ...
, and possibly
Mali Mali (; ), officially the Republic of Mali,, , ff, 𞤈𞤫𞤲𞥆𞤣𞤢𞥄𞤲𞤣𞤭 𞤃𞤢𞥄𞤤𞤭, Renndaandi Maali, italics=no, ar, جمهورية مالي, Jumhūriyyāt Mālī is a landlocked country in West Africa. Ma ...
. Its natural
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
s are subtropical or tropical moist lowland
forest A forest is an area of land dominated by trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing, and ecological function. The United Nations' ...
s, moist
savanna A savanna or savannah is a mixed woodland- grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground ...
,
river A river is a natural flowing watercourse, usually freshwater, flowing towards an ocean, sea, lake or another river. In some cases, a river flows into the ground and becomes dry at the end of its course without reaching another body of ...
s, intermittent rivers,
swamp A swamp is a forested wetland.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p. Swamps are considered to be transition zones because both land and water play a role in ...
s, freshwater
lake A lake is an area filled with water, localized in a basin, surrounded by land, and distinct from any river or other outlet that serves to feed or drain the lake. Lakes lie on land and are not part of the ocean, although, like the much large ...
s, intermittent freshwater lakes, freshwater
marsh A marsh is a wetland that is dominated by herbaceous rather than woody plant species.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p Marshes can often be found ...
es, intermittent freshwater marshes, rural gardens, heavily degraded former forests, water storage areas,
pond A pond is an area filled with water, either natural or artificial, that is smaller than a lake. Defining them to be less than in area, less than deep, and with less than 30% emergent vegetation helps in distinguishing their ecology from tha ...
s, aquaculture ponds, and
canal Canals or artificial waterways are waterways or engineered channels built for drainage management (e.g. flood control and irrigation) or for conveyancing water transport vehicles (e.g. water taxi). They carry free, calm surface f ...
s and ditches.


Description

The western clawed frog is a medium-sized species with a somewhat flattened body and a snout-vent length of , females being larger than males. The eyes are bulging and situated high on the head and there is a short
tentacle In zoology, a tentacle is a flexible, mobile, and elongated organ present in some species of animals, most of them invertebrates. In animal anatomy, tentacles usually occur in one or more pairs. Anatomically, the tentacles of animals work main ...
just below each eye. A row of unpigmented dermal tubercles runs along the flank from just behind the eye, and are thought to represent a
lateral line The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelial ...
organ. The limbs are short and plump, and the fully webbed feet have horny claws. The skin is finely granular. The dorsal surface varies from pale to dark brown and has small grey and black spots. The ventral surface is dull white or yellowish with some dark mottling.


Distribution and habitat

The western clawed frog is an aquatic species and is found in the West African rainforest belt with a range stretching from Senegal to Cameroon and eastern Zaire. It is generally considered a forest-dwelling species and inhabits slow-moving streams, but it is also found in pools and temporary ponds in the northern Guinea and Sudan
savanna A savanna or savannah is a mixed woodland- grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground ...
s.


Biology

In the dry season, this frog lives in shallow streams and hides under tree roots, under flat stones, or in holes in the riverbank. It feeds primarily on earthworms, insect larvae and tadpoles. When the rainy season starts it migrates across the forest floor at night to find temporary pools. Spawning may take place in large pools with much vegetation, but tadpoles are also sometimes found in muddy pools with no vegetation. Single eggs may be attached to plants or they may float. The
tadpole A tadpole is the larval stage in the biological life cycle of an amphibian. Most tadpoles are fully aquatic, though some species of amphibians have tadpoles that are terrestrial. Tadpoles have some fish-like features that may not be found ...
s have broad mouths and no jaws, but have long tentacles on their upper lips. The ventral fins of their tails are broader than the dorsal ones. Their body colour is generally orange and the tail transparent but in darker locations the tail may be blackish. The tadpoles feed by filtering
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
from the water. In large water bodies, they may form dense swarms.
Metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some inse ...
takes place when the tadpoles measure about in length.


Sex determination

Sex determination in the vast majority of
amphibian Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbo ...
s is controlled by homomorphic (morphologically indistinguishable)
sex chromosome A sex chromosome (also referred to as an allosome, heterotypical chromosome, gonosome, heterochromosome, or idiochromosome) is a chromosome that differs from an ordinary autosome in form, size, and behavior. The human sex chromosomes, a typical ...
s. As a result of this difficulty in sex chromosome identification, only a relatively small proportion of anuran species that have been
karyotype A karyotype is the general appearance of the complete set of metaphase chromosomes in the cells of a species or in an individual organism, mainly including their sizes, numbers, and shapes. Karyotyping is the process by which a karyotype is disce ...
d have also had their sex chromosomes identified. Of the species in the genus
Xenopus ''Xenopus'' () (Gk., ξενος, ''xenos''=strange, πους, ''pous''=foot, commonly known as the clawed frog) is a genus of highly aquatic frogs native to sub-Saharan Africa. Twenty species are currently described within it. The two best-know ...
, all have homomorphic sex chromosomes. Additionally, the DM-W gene on the W chromosome in some
Xenopus ''Xenopus'' () (Gk., ξενος, ''xenos''=strange, πους, ''pous''=foot, commonly known as the clawed frog) is a genus of highly aquatic frogs native to sub-Saharan Africa. Twenty species are currently described within it. The two best-know ...
species is the only sex-determining gene that has been identified in amphibians. This DM-W gene was first identified in ''X. laevis'', however it is not found in ''X. tropicalis''. Experimentation involving sex-reversed individuals, gynogenesis,
triploids Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contain ...
, and conventional crosses, has determined that ''X. tropicalis'' has three sex chromosomes: Y, W, and Z. These three sex chromosomes produce three different male genotypes, YW, YZ, and ZZ (all are phenotypically identical) and two different female genotypes, ZW, and WW (all are phenotypically identical). As a result, offspring of ''X. tropicalis'' can have sex ratios that differ from the commonly known 1:1 usually found in species with only two different sex chromosomes. For example, offspring resulting from a ZW female and a YZ male will have a sex ratio of 1:3 females to males and offspring resulting from a WW female and a ZZ male will be all female. As a result of this sex determination system, both male and female ''X. tropicalis'' can be either heterogametic or homogametic which is extremely rare in nature. The exact genetic mechanism and the exact alleles underlying this system is not yet known. One possible explanation is that the W chromosome contains a female-determining allele that has function that is not found on the Z chromosome while the Y chromosome contains an allele that acts a negative regulator that is dominant over the female-determining allele on the W chromosome. Although ''X. tropicalis'' does have these three sex chromosomes, the frequency of these three sex chromosomes is not evenly distributed among this species' populations throughout its natural range. The Y chromosome has been identified from two localities in Ghana and in a laboratory strain that originated in Nigeria and the Z chromosome has been confirmed to exist in individuals from western and eastern Ghana. Additionally, all three sex chromosomes have been found to exist together in ''X. tropicalis'' populations in Ghana and potentially elsewhere in its range as well. Additionally, having irregular sex ratios in offspring is generally thought to be disadvantageous so whether or not the existence of three sex chromosomes in ''X. tropicalis'' is evolutionarily stable or an indication that the species is going through a sex chromosome transition (turnover), is still a question. It seems likely that the emergence of the Y chromosome is the most recent event in the evolution of this species' sex chromosomes. It is possible that in the future extinction of the Z chromosome would make it so that the W chromosome transitions into a X chromosome making this species with sex determined by an XY system. It is also possible that if the Y chromosome were to go extinct, this species will have reverted to using an ancestral ZW system.


Status

The
IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
lists the western clawed frog as "
Least Concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. ...
" because it has a wide distribution and is an adaptable species living in a range of habitats, and the population trend seems to be steady.


Use as a genetic model system

:''See also
Xenopus ''Xenopus'' () (Gk., ξενος, ''xenos''=strange, πους, ''pous''=foot, commonly known as the clawed frog) is a genus of highly aquatic frogs native to sub-Saharan Africa. Twenty species are currently described within it. The two best-know ...
- Model organism for biomedical research'' ''Xenopus'' embryos and eggs are a popular model system for a wide range of biomedical research. This animal is widely used because of its powerful combination of experimental tractability and close evolutionary relationship with humans, at least compared to many model organisms.Wallingford, J., Liu, K., and Zheng, Y. 2010.''Current Biology'' vol. 20, p. R263–4 Unlike its sister species ''X. laevis'', ''X. tropicalis'' is diploid and has a short
generation time In population biology and demography, generation time is the average time between two consecutive generations in the lineages of a population. In human populations, generation time typically ranges from 22 to 33 years. Historians sometimes use this ...
, facilitating genetic studies. The complete genome of ''X. tropicalis'' has been sequenced. This species has n=10 chromosomes. ''X. tropicalis'' has three
transferrin Transferrins are glycoproteins found in vertebrates which bind to and consequently mediate the transport of iron (Fe) through blood plasma. They are produced in the liver and contain binding sites for two Fe3+ ions. Human transferrin is encod ...
genes, all of which are close
ortholog Sequence homology is the biological homology between DNA, RNA, or protein sequences, defined in terms of shared ancestry in the evolutionary history of life. Two segments of DNA can have shared ancestry because of three phenomena: either a s ...
s of other vertebrates. They are relatively far from non-vertebrate chordates, and widely divergent from
protostome Protostomia () is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryonic development. This nature has since been discovered to be extremely variable among Protostomia's me ...
orthologs.


Online Model Organism Database

Xenbase is the Model Organism Database (MOD) for both ''
Xenopus laevis The African clawed frog (''Xenopus laevis'', also known as the xenopus, African clawed toad, African claw-toed frog or the ''platanna'') is a species of African aquatic frog of the family Pipidae. Its name is derived from the three short claws o ...
'' and ''Xenopus tropicalis''.


References


External links


Xenbase
''Xenopus'' model organism database *View th
xenopus genome
in
Ensembl Ensembl genome database project is a scientific project at the European Bioinformatics Institute, which provides a centralized resource for geneticists, molecular biologists and other researchers studying the genomes of our own species and other v ...
* {{Taxonbar, from=Q2705052 Xenopus Amphibians described in 1864 Taxa named by John Edward Gray Animal models