Hynobius Fossigenus
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''Hynobius fossigenus'' is a species of
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All t ...
in the family
Hynobiidae The Asiatic salamanders (family Hynobiidae) are primitive salamanders found all over Asia, and in European Russia. They are closely related to the giant salamanders (family Cryptobranchidae), with which they form the suborder Cryptobranchoidea. ...
, and is found in some
prefectures A prefecture (from the Latin ''Praefectura'') is an administrative jurisdiction traditionally governed by an appointed prefect. This can be a regional or local government subdivision in various countries, or a subdivision in certain international ...
in Kantō and Chūbu districts in Japan. Like all species in the genus ''
Hynobius ''Hynobius'' is a genus of salamander (Asian salamanders) in the family Hynobiidae, occurring in Japan, Korea, China, Taiwan and Far East Russia. It contains these species: Species Species included (as of March 2021): *'' Hynobius abei'' Sat ...
'', it has
lungs The lungs are the primary organs of the respiratory system in humans and most other animals, including some snails and a small number of fish. In mammals and most other vertebrates, two lungs are located near the backbone on either side ...
, its tail is smaller than its body, and it does not have
claws A claw is a curved, pointed appendage found at the end of a toe or finger in most amniotes (mammals, reptiles, birds). Some invertebrates such as beetles and spiders have somewhat similar fine, hooked structures at the end of the leg or tarsus ...
on its fingertips. However, characteristics such as its purplish coloration with golden spots, its size, and its use of lotic waters to reproduce make this species different from all the others. The individuals breed between December and April in mountain streams surrounded by ''
Cryptomeria japonica ''Cryptomeria'' (literally "hidden parts") is a monotypic genus of conifer in the cypress family Cupressaceae, formerly belonging to the family Taxodiaceae. It includes only one species, ''Cryptomeria japonica'' ( syn. ''Cupressus japonica'' L ...
''
evergreen forests An evergreen forest is a forest made up of evergreen trees. They occur across a wide range of climatic zones, and include trees such as conifers and holly in cold climates, eucalyptus, Live oak, acacias, magnolia, and banksia in more temperate zon ...
. After mating, the females lay their eggs in envelopes, which are thick, transparent, and resistant, where they remain for sixty days, when the embryo fully develops in the egg. After hatching, the
tadpoles 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 in ...
leave the envelope and start living under rocks or at the bottom of streams, completing their metamorphosis within a year. The males become adults around the age of five, while the females become adults around the age of seven.


Taxonomy

The species was described in the scientific journal
PeerJ ''PeerJ'' is an open access peer-reviewed scientific mega journal covering research in the biological and medical sciences. It is published by a company of the same name that was co-founded by CEO Jason Hoyt (formerly at Mendeley) and publisher ...
on June 21, 2018, by researchers Hisanori Okamiya, Hirotaka Sugawara, Masahiro Nagano, and Nikolay A. Poyarkov. It is treated as belonging to the genus ''
Hynobius ''Hynobius'' is a genus of salamander (Asian salamanders) in the family Hynobiidae, occurring in Japan, Korea, China, Taiwan and Far East Russia. It contains these species: Species Species included (as of March 2021): *'' Hynobius abei'' Sat ...
'', since it has all the characteristics that together are unique to it, such as: the presence of
lungs The lungs are the primary organs of the respiratory system in humans and most other animals, including some snails and a small number of fish. In mammals and most other vertebrates, two lungs are located near the backbone on either side ...
, having a tail smaller than the body, existence of clawless fingers, and the absence of a clear, visible line on the dorsal region. Speciation can be determined by its large size, by laying its eggs in lotic waters, by the thickness of its egg sac, by its small head and slender body, and by its coloration. Genetic tests were also made from its
nuclear Nuclear may refer to: Physics Relating to the nucleus of the atom: * Nuclear engineering *Nuclear physics *Nuclear power *Nuclear reactor *Nuclear weapon *Nuclear medicine *Radiation therapy *Nuclear warfare Mathematics *Nuclear space *Nuclear ...
and mitochondrial DNA, as well as from mitochondrial
16S RNA 16 S ribosomal RNA (or 16 S rRNA) is the RNA component of the 30S subunit of a prokaryotic ribosome (SSU rRNA). It binds to the Shine-Dalgarno sequence and provides most of the SSU structure. The genes coding for it are referred to as 16S rR ...
, which attested that it is indeed a new species. The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...
, which was an adult male, was found on March 1, 2018, in a stream on the eastern slope of Mount Hinode in the town of the same name in Japan at an elevation of 679 meters. Also used in the description were 25
paratypes In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype ...
, 17 males and eight females, which were collected during the breeding season in the surrounding regions of the town where the holotype was found. Since 1986, there were already suspicions that the species existed, when researchers Ikebe, Yamamoto, and Kohno, when analyzing different populations of '' Hynobius kimurae'', noticed a certain difference in the karyotype between individuals, in addition to a
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
proximity of this species with ''H. boulengeri''. Later, in another study in 1991, Ikebe and Kohno detected that this heterogeneity was located in the Kantō district, in addition to noticing several distinctions between the groups, such as the number of eggs per clutch and the biology of the
tadpoles 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 in ...
. Despite all these dissimilarities, the species was not described at that time, and the data was attributed to mere divergences between distinct population groups. The species' specific epithet is an agglutination of the Latin words ''fossa'', meaning abyss, and ''genus'', meaning born in, forming an adjective that loosely translates as one born in the abyss, a reference to the fact that it is found near a fault between the
North American Plate The North American Plate is a tectonic plate covering most of North America, Cuba, the Bahamas, extreme northeastern Asia, and parts of Iceland and the Azores. With an area of , it is the Earth's second largest tectonic plate, behind the Pacif ...
and the
Eurasian Plate The Eurasian Plate is a tectonic plate that includes most of the continent of Eurasia (a landmass consisting of the traditional continents of Europe and Asia), with the notable exceptions of the Indian subcontinent, the Arabian subcontinent and ...
.


Distribution and conservation

The species can be found in several
prefectures A prefecture (from the Latin ''Praefectura'') is an administrative jurisdiction traditionally governed by an appointed prefect. This can be a regional or local government subdivision in various countries, or a subdivision in certain international ...
in Japan, such as
Gunma is a prefecture of Japan located in the Kantō region of Honshu. Gunma Prefecture has a population of 1,937,626 (1 October 2019) and has a geographic area of 6,362 km2 (2,456 sq mi). Gunma Prefecture borders Niigata Prefecture and Fukushima P ...
, Saitama,
Tokyo Tokyo (; ja, 東京, , ), officially the Tokyo Metropolis ( ja, 東京都, label=none, ), is the capital and List of cities in Japan, largest city of Japan. Formerly known as Edo, its metropolitan area () is the most populous in the world, ...
,
Kanagawa is a prefecture of Japan located in the Kantō region of Honshu. Kanagawa Prefecture is the second-most populous prefecture of Japan at 9,221,129 (1 April 2022) and third-densest at . Its geographic area of makes it fifth-smallest. Kanagaw ...
, Yamanashi,
Shizuoka Shizuoka can refer to: * Shizuoka Prefecture, a Japanese prefecture * Shizuoka (city), the capital city of Shizuoka Prefecture * Shizuoka Airport * Shizuoka Domain, the name from 1868 to 1871 for Sunpu Domain, a predecessor of Shizuoka Prefecture ...
,
Nagano Nagano may refer to: Places * Nagano Prefecture, a prefecture in Japan ** Nagano (city), the capital city of the same prefecture *** Nagano 1998, the 1998 Winter Olympics *** Nagano Olympic Stadium, a baseball stadium in Nagano *** Nagano Universi ...
, and
Aichi is a prefecture of Japan located in the Chūbu region of Honshū. Aichi Prefecture has a population of 7,552,873 () and a geographic area of with a population density of . Aichi Prefecture borders Mie Prefecture to the west, Gifu Prefecture ...
. Many of these populations are isolated from each other by landforms such as massifs. It is present between the altitude range of 300 and 1 100 meters, but is most common between 400 and 900 meters. It is usually close to lotic waters , such as streams, where they lay their eggs. Although the species is not protected by any law or environmental park, with the exception of the Gunma prefecture, it has a wide distribution, and only a few isolated populations may be threatened by anthropogenic actions or
habitat loss Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
, making it a
least-concern species 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. T ...
(LC). It has not yet been assessed by the
International Union for Conservation of Nature 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 nat ...
(IUCN).


Speciation

Through genetic tests performed on the species, it was possible to determine that speciation occurred from the
adaptive radiation In evolutionary biology, adaptive radiation is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic int ...
of the
specific Specific may refer to: * Specificity (disambiguation) * Specific, a cure or therapy for a specific illness Law * Specific deterrence, focussed on an individual * Specific finding, intermediate verdict used by a jury in determining the fina ...
''H. kimurae-H. boulengeri'' complex, which occurred seven million years ago, at the end of the
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
. Such radiation was caused by the appearance of mountains of volcanic origin, which separated the complex into three different groups: the western, which gave rise to ''H. kimurae'', the central, ''H. boulengeri'', and the eastern, ''H. kimurae''. These populations, being reproductively isolated, and due to
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
, began to diverge and became separate species.


Description

Its average length varies between 66 and 82.5 millimeters, of which only the head, which is oval and broad, accounts for 25%. Its body is slender and cylindrical, with a narrow
thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the cre ...
. The skin of its belly and back is smooth, with microscopic glands scattered all over the body. Its cloaca is slightly swollen, not visible from the ventral or lateral point of view, and is open longitudinally in the shape of a cross, with prominent edges and a reduced sex
tubercle In anatomy, a tubercle (literally 'small tuber', Latin for 'lump') is any round nodule, small eminence, or warty outgrowth found on external or internal organs of a plant or an animal. In plants A tubercle is generally a wart-like projection ...
. The tail is long, but smaller than the body, ranging in shape from cylindrical to oval, and thicker near the base. Its tongue is broad, elliptical, and convex, and is attached to the lower part of the mouth by the central part. The muzzle is wide, short, and rounded, with small, round nostrils pointed to the side. Its eyes are large and the upper eyelid is well developed. They have prominent, swollen parotoid glands, which extend in the region between the
lower jaw In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
and the
throat In vertebrate anatomy, the throat is the front part of the neck, internally positioned in front of the vertebrae. It contains the pharynx and larynx. An important section of it is the epiglottis, separating the esophagus from the trachea (windpip ...
fold. Their
dental arch The dental arches are the two arches (crescent arrangements) of teeth, one on each jaw, that together constitute the dentition. In humans and many other species; the superior (maxillary or upper) dental arch is a little larger than the inferior ...
is composed of long, wide, oblique rows of usually 56 U-shaped
vomerine teeth The vomer (; lat, vomer, lit=ploughshare) is one of the unpaired facial bones of the skull. It is located in the midsagittal line, and articulates with the sphenoid, the ethmoid, the left and right palatine bones, and the left and right maxill ...
. The coloration of its back varies from brownish-purple to faded black, and it has several small irregularly shaped spots distributed throughout the body, which can be golden yellow or golden orange, the amount of which can vary among individuals. Its legs have a more grayish coloration than the rest of the body. The main
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...
is the size of the males, which is usually smaller than the females, and the tail, which is usually higher. During the reproductive period, the cloaca of males is usually more protuberant, the opposite of what happens with females, in which the organ is less prominent. During reproduction, in turn, the female's body tends to become more swollen. It can be differentiated from all
Taiwanese Taiwanese may refer to: * Taiwanese language, another name for Taiwanese Hokkien * Something from or related to Taiwan ( Formosa) * Taiwanese aborigines, the indigenous people of Taiwan * Han Taiwanese, the Han people of Taiwan * Taiwanese people, ...
congeneric species, such as ''H. formosanus'' and ''H. glacialis'' by its coloration and large size, as they never exceed 69 millimeters. It can be differentiated from all the lytic species of its
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
by the fact that its egg sac is sturdier and has more folds, and has a violet-fluorescent coloration. ''H. katoi'', a sympatric species, can be differentiated by being smaller (60 millimeters) and having fewer teeth, a total of 39. Of the species of its specific complex, it can be differentiated from ''H. boulengeri'' by having a completely uniform bluish-black coloration on its back, and from ''H. kimurae'', the phylogenetically closest species, by being larger than the latter, which has 74 millimeters, a longer tail, and by its coloration.


Reproduction

The species usually breeds in the
spring Spring(s) may refer to: Common uses * Spring (season) Spring, also known as springtime, is one of the four temperate seasons, succeeding winter and preceding summer. There are various technical definitions of spring, but local usage of ...
of small mountain streams, with temperatures never exceeding 20°C. The preferred places are those less than one and a half meters wide and twenty to thirty centimeters deep, located in
evergreen In botany, an evergreen is a plant which has foliage that remains green and functional through more than one growing season. This also pertains to plants that retain their foliage only in warm climates, and contrasts with deciduous plants, whic ...
''
Cryptomeria japonica ''Cryptomeria'' (literally "hidden parts") is a monotypic genus of conifer in the cypress family Cupressaceae, formerly belonging to the family Taxodiaceae. It includes only one species, ''Cryptomeria japonica'' ( syn. ''Cupressus japonica'' L ...
'' or mixed forests. Adults begin searching for such sites in November, with the reproductive period lasting from December to April. Mating occurs out of the water, in burrows under rocks and stones. After that, the females lay their eggs in thick, resistant, transparent envelopes, which are attached to the rocks in the stream or waterfalls, where the water is usually between 5.5 and 6.5°C. Soon after, the female leaves the site, and the male usually stays for a while in the surrounding region where oviposition occurred. The eggs usually develop for sixty days, and after that time they hatch and the tadpoles come out of the envelope. Outside, they usually feed on
amphipods Amphipoda is an order of malacostracan crustaceans with no carapace and generally with laterally compressed bodies. Amphipods range in size from and are mostly detritivores or scavengers. There are more than 9,900 amphipod species so far describ ...
and larvae of tricoptera and ephemeroptera. There are reports of cannibalism, in which the older ones feed on the younger ones. They are easily found under rocks and fallen leaves or in the deepest part of streams. The metamorphosis lasts about a year, and when it is over, the imagos start feeding on spiders, insects, and worms. Males take a minimum of five years to become adults, while females take a little longer, with a minimum of seven years.


References

* * {{Taxonbar, from=Q55778985 fossigenus Amphibians of Japan Amphibians described in 2018 Species described in 2018