Japanese fire-bellied newt

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Japanese fire-bellied newt
Temporal range: 13.75–0 Ma Middle Miocene – Present
Cynops pyrrhogaster sasayamae female.jpg
Scientific classification edit
Kingdom: Animalia
Phylum: Chordata
Class: Amphibia
Order: Urodela
Family: Salamandridae
Genus: Cynops
Species:
C. pyrrhogaster
Binomial name
Cynops pyrrhogaster
(Boie, 1826)
Fire-bellied newt range.png
Synonyms

see text

The Japanese fire-bellied newt or Japanese fire-bellied salamander (Cynops pyrrhogaster) is a species of newt endemic to Japan. While officially a single species, it appears in four distinct varieties. The skin on its upper body is dark, while its lower regions are bright red, although coloration varies with age, genetics, and region. They contain high levels of tetrodotoxin, a neurotoxin, accumulated mainly from diet. They are found on many Japanese islands, especially the major ones. Their habitat includes both natural and artificial bodies of water, as well as forests and grasslands. They breed from spring to the beginning of summer, both sexes producing pheromones when ready. Eggs are laid separately, hatching after about 3 weeks.

They grow from larval to juvenile form in between five and six months. Juveniles eat soil-dwelling prey, while adults eat a wide variety of insects, tadpoles, and potentially the eggs of their own species. They have multiple adaptations to avoid predators, although the type they use depends on where they live. Multiple aspects of their biology have been studied, including their ability to regrow missing body parts. Their population is declining, and they face threats from disease and the pet trade. They can be successfully kept in captivity.

Etymology and taxonomy

The species was first described by Heinrich Boie in 1826 as Molga pyrrhogaster,[note 1] in his work Kenteekenen van eenige Japansche Amphibiën. He compared it to the smooth newt, saying he would have mistaken the former species for the latter, had he not known it was from Japan. None of the specimens he studied were fully mature.[2] It was moved to the genus Cynops in 1838 by Johann Jakob von Tschudi, as Cynops subcristatus.[3] In 1850, it was given its current binomial name of Cynops pyrrhogaster by John Edward Gray.[4] The Integrated Taxonomic Information System recognizes sixteen total synonyms.[5] Common names of Cynops pyrrhogaster include the Japanese fire-bellied newt,[1] red-bellied newt,[6] and Japanese fire-bellied salamander.[7]

Traditionally, six races were recognized (Tohoku, Kanto, Atsumi, intermediate, Sasayama, and Hiroshima), however later molecular analysis supported their division into four clades instead.[8] In particular, the validity of the Sasayama and intermediate races has never been proven, with one study finding no behavioral differences between the two supposed forms.[9]

A study of mitochondrial DNA in 2001 indicated that its supposed fellow members of Cynops, C. cyanurus and C. wolterstorffi, may in fact belong to a different genus.[10]

Evolution

C. pyrrhogaster diverged from its close relative, C. ensicauda, about 13.75 million years ago (during the Middle Miocene). The common ancestor of the two species would have lived on an area of the Eurasian mainland which is today the East China Sea and the central Ryukyu Islands. In that time, the land that would become the Japanese islands had a subtropical climate, which may have caused the Japanese fire-bellied newt's ancestors to migrate northward for desirable habitat. As time progressed, C. pyrrhogaster split into four clades — northern, southern, western, and central. The northern diverged first, at around 9.68 million years ago, then the central (around 8.23 MYA), then finally the southern and western (around 4.05 MYA). The ranges of all but the southern clade declined during Last Glacial Period, but expanded again afterwards. The study that identified them concluded that the four clades represent separate taxonomic units, although their exact relationship is unclear.[8]

Hybrids

The ranges of the central and western varieties meet in Chūgoku in western Japan to form a hybrid zone. The central type has begun to move west, which has caused the hybrid zone to shift. It is expected to eventually cause the genome of the western form to be lost to increasing hybridization.[11]

Description

Japanese fire-bellied newt

The skin is dark brown, approaching black, and covered in wartlike bumps. Its underbelly and the underside of its tail are bright red, with black spots.[2] Specimens from smaller islands tend to have more red on their ventral regions than those from larger islands, sometimes with extremely small spots or none at all.[12] Smaller juveniles have creamy coloration instead of red, although most larger juveniles have red present.[13] Males can be distinguished from females by their flat, wide tails and swelling around the ventral region.[14] An entirely red variant exists, believed to be inherited and recessive.[15]

The vomeropalatine teeth are arranged in two series. The tongue is relatively small, half the width of the mouth. The nostrils are positioned anteriorly, closer to each other than to the eyes, hardly visible when viewed from above. The toes of males are longer than those of females, although the females themselves are longer. The tail is tightly compressed, with fins on both the top and bottom. A smooth ridge runs from their nape to their tail.[16] It is 8 to 15 centimetres (3.1 to 5.9 in) long.[7] Eggs are 2.1 to 2.3 millimetres (0.083 to 0.091 in) in size.[14]

Toxin

Bright red ventral area

Wild Japanese fire-bellied newts contain high levels of the neurotoxin tetrodotoxin (TTX). Experimentation has found that this is almost entirely from diet. When raised in captivity with no source of TTX, 36- to 70-week-old juveniles did not contain detectable levels, while wild specimens from the same original habitat had high toxicity. However, in younger captive-reared newts, some TTX was still detected, which was inferred to have been transferred to their eggs from the adult females.[17]

In a follow-up experiment by the same team, captive-reared newts were given food containing the neurotoxin. They readily consumed TTX-laced bloodworms when offered, not showing any symptoms after ingesting the poison.[18]

Atsumi-Chita variant

A variety believed to be found exclusively on the Atsumi Peninsula was thought to be extinct, however it was later revealed that newts on the Chita Peninsula shared many characteristics and very likely belong to the same race. Both groups share a preference for cooler temperature, smooth and soft bodies, pale dorsal regions, and yellowish undersides. Even if still alive, this race is highly threatened and requires immediate protection, or it will soon be wiped out.[19]

Distribution and habitat

Cynops pyrrhogaster is endemic to Japan, found on numerous islands in the archipelago, including Honshu, Shikoku, and Kyushu.[1] Primarily, it dwells on the major islands, while its relative, C. ensicauda, is found in the Ryuku Islands. It has the northernmost range of any Cynops species.[8] There is also an introduced population on Hachijojima Island, believed to be descended from individuals from Shikoku.[20] It has been recorded in the United States three times, in the states of Florida and Massachusetts. Every instance was either an escape or deliberate release, and no populations have been established.[7]

They occur at elevations of 30 metres (98 ft) to 2,020 metres (6,630 ft). Ecosystems they are found in include forests, grasslands, shrublands, wetlands, lakes, marshes, and cultivated environments. They can also dwell in humanmade bodies of water, such as aquaculture ponds.[1]

Behavior and ecology

Pair of C. pyrrhogaster specimens

Reproduction and life cycle

Reproduction

Breeding occurs in paddy fields, ponds, brooks, pools, and streams. Females accept male courtship behavior from spring to early summer.[21]

Males and females both produce peptide pheromones to attract the opposite sex when ready to mate. Males produce a type known as sodefrin, while females have their own variety, named "imorin" by its discoverers (from the Japanese word imo for "beloved woman" and rin from sodefrin). These are released from the cloaca, and were the first peptide pheromone to be identified in a vertebrate and first to be identified in a female vertebrate, respectively.[6][22]

Courtship begins when the male approaches the female, sniffing its sides or cloaca. The male then brings its tail to the female and rapidly vibrates it. The female responds by pushing the male's neck with its snout. At this point, the male slowly moves away, undulating its tail, while the female follows, touching the tail with its snout when close enough. The male then deposits two to four spermatophores, one at a time, moving several centimeters after each, which the female attempts to pick up with its cloaca, sometimes unsuccessfully.[23]

Females lay eggs separately on underwater objects, such as leaves and submerged grass roots, fertilized one-by-one via the spermatophores they carry. They can lay up to 40 eggs in one session, and 100 to 400 eggs in a full breeding season.[23]

Life cycle

Unlike larvae, older newts cannot stay underwater indefinitely

The young hatch from their eggs after about 3 weeks, as swimming, gilled larvae, with dorsal tailfins. In captive settings, they are known to readily eat mosquito larvae, brine shrimp, earthworms, and each other. They grow around 3 centimetres (1.2 in) in the first three months of their lives. They undergo metamorphosis at between five and six months (immediately before which, they stop eating), losing their gills and fins, and becoming juveniles. Juveniles cannot remain submerged in water like larvae or they will drown.[24][25]

Newts at lower altitudes mature faster than those at higher ones, and male newts of these populations tend to live longer after reaching maturity. However, their fully-grown size is not as large as that of lowland newts. Wild individuals as old as twenty-three have been found.[21]

Diet

Juveniles often consume soil-dwelling Collembola and Acari species.[13] Adults at one particular sub-alpine moor in the Azuma Mountains of the Fukushima Prefecture were found to take both live prey and animals that were already dead. They consume many insect varieties, such as members of Odonata (larvae have been found whole in newt stomachs, while only pieces of adults have been discovered), Brachycera, Hymenoptera, and Coleoptera. They also eat Rhacophorus arboreus tadpoles and the eggs of their own kind. The makeup of their diet varies seasonally and from year-to-year, suggesting changes in the small animals around the ponds that they dwell in.[14] Similar results were found at a pond on the campus of Tokyo Metropolitan University in Hachiōji, Tokyo, with newt stomachs containing insects from many different orders, and again, the eggs of conspecifics. Again, frog tadpoles were eaten, although these belonged to the species Rhacophorus schlegelii.[26]

Predators

Newts in Mainland Japan have different antipredator behavior than newts on smaller ones. Individuals on smaller islands (for instance, Fukue Island) generally utilize a maneuver called the unken reflex, where they expose their bright red underbelly to attackers. As their main predators are birds, which are capable of distinguishing the color red, this technique is effective. However, in Mainland Japan, the newts must also avoid mammalian predators, which cannot distinguish colors as well as avian hunters. This leads these populations to use the maneuver less, as it can result in death if attempted.[12]

Against snakes, newts from Fukue Island tend to perform tail-wagging displays (which are designed to bring a predator's attention to their replaceable tail rather than their more valuable head), while those from Nagasaki Prefecture, in Mainland Japan tend to simply flee. Snakes are present in both areas, with one example of a potential serpentine predator being Gloydius blomhoffii. This is likely because those from the mainland are adapted to escape from mammalian hunters, which are less likely to be repelled by such a display.[27]

Research

Moliton

Japanese fire-bellied newts produce motilin, a peptide that stimulates gastrointestinal contractions in mammals. It is created in the upper small intestine and pancreas (the discovery of the latter was the first time pancreatic motilin had been observed). Their pancreas also produces insulin.[28]

Regeneration

This species, as well as other Urodele amphibians, is capable of regrowing missing body parts, including limbs with functional joints and the lower jaw.[29][30] When this process occurs, the regenerated tissue tends to mirror intact tissue in form.[29] It is also able to regrow missing lenses, taking 30 days to do so as a larva and 80 days as an adult (the difference in time is purely due to the size of the eye, and regenerative ability does not change).[31]

Metamorphosis inhibition

While a highly useful model organism, they become more difficult to care for in lab settings after metamorphosis. An experiment supported by the Japan Society for the Promotion of Science found that thiourea (TU) can prevent this process from occurring, allowing the animals to stay in their pre-metamorphosis form for as long as two years, while still capable of metamorphosizing when removed from the TU solution. This did not have any impact on their regeneration capabilities.[24]

Conservation

Group at Ikenokouchi Wetland in Tsuruga, Fukui Prefecture

The International Union for the Conservation of Nature (IUCN) has ranked it as near threatened. This assessment was made in 2020,[1] a shift from 2004, when it was rated least-concern.[32] It successfully breeds in captivity at Australian zoos.[1]

Threats

One major threat that C. pyrrhogaster faces is collection for the pet trade. The IUCN states that this trade needs to be ended "immediately". Their population is decreasing, particularly near areas of human habitation.[1]

Disease

Japanese fire-bellied newts with mysterious skin lesions at Lake Biwa in Japan's Shiga Prefecture were found to be suffering from infections caused by a single-celled eukaryote in the order Dermocystida. The leisons contained cysts, which were filled with spores. Nearly all the lesions were external, although one was found on the liver. Globally, diseases are one of the causes for declining amphibian populations. There is concern that this affliction could spread to other nearby species, including Zhangixalus arboreus and Hynobius vandenburghi).[33]

In captivity

Japanese fire belly newt crawling

C. pyrrhogaster can be kept in captivity. DVM Lianne McLeod described them as "low-maintenance". She wrote that that captive newts enjoy bloodworms, brine shrimp, glass shrimp, daphnia, and, for larger individuals, guppies. She also stressed the need to maintain a clean tank, as disease and death can result from dirty gravel or water.[34]

Notes

  1. ^ Later authors use the spelling Molge.

References

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