The Smiling Spirits of the Mountains: Salamanders and Their Remarkable Fatherhood

When people think of animals that embody strong family bonds, mammals and birds usually come to mind first. Kangaroos in Australia, the cats and dogs that share our homes, and many familiar birds are often celebrated for their parental devotion. Yet few people realize that among amphibians there are also species whose lives are deeply shaped by parental care. The salamanders introduced here belong to one such group—animals whose reproductive behavior reveals an extraordinary form of paternal devotion.

Despite the word "fish" appearing in their Chinese name, salamanders are not fish at all. They are tailed amphibians belonging to the suborder Cryptobranchoidea within the order Caudata. This suborder includes two families: Cryptobranchidae and Hynobiidae. Members of Cryptobranchidae include the giant salamanders of the genus Andrias, often referred to in Chinese as "baby fish." In Taiwan, however, the native mountain salamanders belong to the family Hynobiidae.

Five species of hynobiid salamanders occur in Taiwan: Hynobius fuca (Guangwu salamander), Hynobius glacialis (Nanhu salamander), Hynobius formosanus (Taiwan salamander), Hynobius sonani (Sonan's salamander), and Hynobius arisanensis (Alishan salamander). All five inhabit high-mountain environments, typically living at elevations between 2000 and 3000 meters.

Why are salamanders described as animals full of paternal devotion? The answer lies in their unusual reproductive behavior. Like most amphibians, salamanders undergo two major life stages during their life cycle: an aquatic larval stage and a terrestrial adult stage. Their breeding season runs from September to January. During this period, male and female salamanders leave their usual habitats and travel to nearby streams to find mates.

Unlike frogs, salamanders do not attract partners through vocal calls. Instead, males perform a distinctive courtship dance to display themselves to females. If rival males appear during the courtship process, competition can escalate into physical confrontation as they struggle for the opportunity to mate.

Figure 1. A male Hynobius arisanensis caring for its eggs, Courtesy of 山椒魚國際保育研討會

After this intense courtship period, a female chooses the male she prefers. The pair then move upstream together toward deeper subterranean water channels beneath the streambed. Compared with the fast-flowing surface current, these underground waters are calmer and better concealed. Once they reach the breeding site, the pair coil around each other in a behavior sometimes described as pseudo-copulation. During this process, the male gently bites the female's neck to anchor the pair together, releasing sperm as the female lays her eggs so that fertilization occurs externally.

Figure 2. Salamander eggs at the neurula stage, Courtesy of 山椒魚國際保育研討會

Unlike mammals, where mothers typically provide most parental care, salamander reproduction places the entire responsibility on the father. After fertilization, the male remains beside the egg sacs, wrapping himself around them to guard them from hungry predators and even cannibalistic members of his own species. From that moment onward, the salamander father becomes a tireless guardian, standing watch day and night until the larvae finally hatch.

Figure 3. A male Hynobius arisanensis arranging the egg sacs, Courtesy of 山椒魚國際保育研討會

The newly hatched larvae differ noticeably from frog tadpoles. Salamander larvae possess distinctive external gills that protrude from the sides of the head. Their appearance is reminiscent of the axolotl (Ambystoma mexicanum), another member of the order Caudata. Soon after hatching, the young larvae leave their father's protection and drift along underground water channels until they reach nearby streams, beginning their lives independently.

Over the following three months, the larvae gradually undergo transformation. Their forelimbs appear first, followed by the hind limbs. Eventually the external gills are lost, and the young salamanders take on the same body form as their parents, entering the terrestrial phase of their life cycle.

Figure 4. A slightly exhausted male Hynobius arisanensis, Courtesy of 山椒魚國際保育研討會

Figure 6. A salamander larva retaining its external gills, Courtesy of 山椒魚國際保育研討會

Figure 7. A salamander larva that has just completed metamorphosis and emerged onto land, Courtesy of 山椒魚國際保育研討會

Despite their charm and remarkable biology, these animals now face serious threats. As global temperatures continue to rise, high-mountain streams are gradually warming. This warming compresses the narrow environmental range in which salamanders can survive. In addition, the rapid expansion of mountain tourism has begun to damage their habitats. Increasing numbers of visitors travel into alpine areas, leaving behind waste and garbage among the rocky slopes. Such disturbances degrade the fragile environments salamanders depend upon, leaving many individuals without suitable homes.

Figure 8. An eco-friendly conference bag printed with a salamander larva, Courtesy of 山椒魚國際保育研討會

Fortunately, growing environmental awareness has inspired many people to take action. Mountain clean-up campaigns have become increasingly common, and researchers are also working to establish artificial breeding habitats that can support salamander populations while natural ecosystems recover. The situation for these gentle mountain spirits remains precarious, but with more people joining conservation efforts, there is hope that these smiling creatures will continue to inhabit these mountains for generations to come.

All photographs above were reproduced by the author during academic conferences.

Author: Rodrigo

Reference:

朱有田，（2022），高山型國家公園山椒魚分佈棲地、遺傳結構與生物學調查，雪霸國家公園管理處（2023）山椒魚國際保育研討會會議手冊，內政部國家公園管理署