Red Hills Salamander
(Phaeognathus hubrichti)
The Red Hills salamander is a large, burrowing species that grows to a length of 250 mm. It was discovered in 1960 and seldom emerges completely from its underground refuge, hunting for invertebrate prey at night at the entrance of its retreat. It lives in steep slopes along damp ravines covered by hardwood forest. Around 60% of its habitat is owned or leased by paper companies that are degrading and destroying this species’ habitat by timber harvest and pine plantations. Six Habitat Conservation Plans for populations of Red Hills salamanders (covering approximately 25,169 hectares) have been approved by the U.S. Fish and Wildlife Service in cooperation with timber companies in southern Alabama.
Urgent Conservation Actions
Maintain conservation partnership with paper/timber companies to selectively conserve occupied habitat along steep slopes; prohibit collection of this species.
South-central Alabama, U.S.A.
Before 1960, the Red Hills Salamander was unknown to science. It was discovered by land snail expert, Leslie Hubricht, in Butler County in 1960 when he was out searching for snails. Instead, he found a large brown salamander. It was later determined by Dr. Richard Highton of the University of Maryland that this salamander was a new species, the first discovered in the United States since 1939. Dr. Highton named the salamander Phaeognathus hubrichti in 1961. Phaeognathus means "dark jawed" and hubrichti is given to honour Leslie Hubricht. The discovery of this species in the Coastal Plain is not surprising since the center of the distribution of the Plethodontinae subfamily is the Appalachian Mountain region.

After its discovery, this species was not collected again until 1963, due to its secretive lifestyle and the fact that it is difficult to catch because it tends to dart back into its burrow. Techniques were subsequently developed to increase the rate of capture, including locating salamanders in their burrow entrances and then driving a pick behind the salamander to prevent its escape down the burrow, and "fishing" for the species in front of its burrow using a small fishhook baited with a cricket or spider. Due to its listing as a federally threatened species, collection of Red Hills salamanders is now prohibited.

The Red Hills salamander has been named the State Amphibian of Alabama.
Evolutionary Distinctiveness
Order: Caudata
Family: Plethodontidae
The Plethodontidae is by far the largest family of salamanders, comprising nearly 70% of all living species. In total there are 378 known plethodontids divided between four subfamilies and 24 genera. The plethodontids are united by the fact that they do not possess lungs and breathe entirely through their skin and mouth lining. They are often referred to as the lungless salamanders, although they are thought to have evolved from highly aquatic, lunged ancestors in the streams of the Appalachian Mountains in eastern North America. The earliest plethodontids were hypothesised to have lost their lungs because individuals with reduced, or absent, lungs were less likely to float away in the swift mountain streams where they lived. The vast majority of other salamanders possess lungs, so this makes the lungless salamanders an unusual and fascinating group of animals.

They are thought to have diverged from all other amphibian species 145 million years ago at the boundary between the Jurassic and Cretaceous periods. They are as different from all other amphibian lineages as wombats are from whales, evolving at a time when dinosaurs were still abundant. Overall, plethodontids are the most evolutionary advanced salamanders, so it may at first appear odd that they should have lost lungs, which are one of the most basic features of all vertebrates living on land. Lacking lungs and being dependent upon their skin for respiration places a size restriction on these salamanders because large animals have a relatively small surface area of skin compared to their body’s volume, and have greater difficulty in supplying their body tissues with oxygen compared to smaller animals (which have a large surface area to volume ratio). The long, slender form of the lungless salamanders maximises the surface area available for gas exchange, and some species grow to lengths of over 300 mm.

Plethodontid salamanders occupy a great diversity of habitats, ranging from strictly aquatic to strictly terrestrial, exploring niches as diverse as caves, trees, mountain streams, and they are also found burrowing through the earth. Dependence on their skin for breathing places limitations upon where and how lungless salamanders can live. Their skin must be kept moist at all times in order for oxygen to be taken up by the blood in capillaries beneath the skin. This means plethodontids are either confined to humid areas, or must find damp hiding places and only emerge in wet weather, typically at night. The life of a lungless salamander in less humid areas, like Europe and temperate North America, therefore comprises brief periods of activity interspersed with inactive phases that are often very long. They are able to survive the periods of inactivity because they have a very low metabolic rate and low energy requirements. Able to store much of what they eat as fat, they do no need to feed very often.

A further adaptation, present among many species of the lungless salamander subfamilies named “Plethodontinae” (from East and West North America) and “Bolitoglossinae” (from tropical Central and South America), is “direct development”. This is a method of amphibian development where the larval stage (e.g. the tadpole stage in a frog’s life history) has been eliminated. Early development takes place in eggs, which may be laid in moist places away from water, and the young hatch out as miniature adults. The well known amphibian metamorphosis, most commonly appreciated in the transition from tadpole to adult frog, does not occur outside of the egg. This mean that certain lungless salamanders in these two subfamilies may live away from water bodies, allowing them to expand their ranges to new areas.

The history and characteristics of the lungless salamanders go some way to explaining their range. They are mostly found in the New World, where they are widely distributed in eastern and western North America, as well as Central and South America. However, continental drift over millions of years has also brought them to the Old World, where they are found in parts Europe (e.g. Sardinia) and Korea. The existence of the Korean crevice salamander was unknown until 2005, when its discovery was a shock to science, indicating a long history of lungless salamanders in Asia. This is the only known species is Asia, suggesting that the rate of species generation in this part of the world is very low, especially compared the the huge radiation of lungless salamander species in the New World.

The Red Hills salamander is one of 93 species that constitutes the Plethodontinae subfamily of East and Northwest America. This species is the only member of its genus (Phaeognathus) and is most closely related to a group of 19 species in the genus Desmognathus, the “dusky” salamanders. The lineage of the Red Hills salamander is thought to have originate 40 millions years ago, which is 5 millions years before the origin of the common ancestor of monkeys and humans, and also the origin dusky salamander lineage. Red Hills salamanders are more elongated than the dusky salamanders in their sister taxon. This is due to an increased number of vertebrae or back bones in the body of this species – most dusky salamanders have 15 vertebrae but the Red Hills salamander has 22. Another unusual feature of this species is that it is one of only three species in the Desmognathus+Phaeognathus group (the so-called desmognathine salamanders) that has direct development of its eggs.

The Red Hills salamander, like all lungless salamanders in the Plethodontidae subfamily, possesses a slender body, long tail and prominent eyes. A distinctive feature of the plethodontid family is a narrow groove (the nasolabial groove) running from each nostril to the upper lip: its function is to carry waterborne odours from the ground into the nasal cavity. Another curious trait of the lungless salamanders are mental (from the Latin “mentum”, meaning chin) glands. These are modified mucus glands and release pheromones, which are chemicals produced by an animal to influence the behaviour of other members if its species, often with regard to breeding receptivity. During amplexus (the mating embrace), the male clasps the female (with both his arms and legs) and rubs pheromones across the female’s snout. Mental glands are sometimes visible in males as raised bumps below their lower lip.

Lungless salamanders are very small to medium in size, usually measuring between 25 to 250 mm from the tip of the nose to the end of the tail, which salamanders retain throughout their life. The Red Hills salamander is one of the largest species of plethodontid, reaching a maximum total length of 250 mm, about half of which is accounted for by a long, tapering tail. It is described as the largets species of woodland salamander. They have well-developed “costal” grooves (successive vertical grooves in the skin along the sides of the body), generally numbering between 20-22. The tail is actually prehensile, which means it can be used to grasp hold of objects, aiding movement. The limbs are slender and short, and this species is dark brown in colour.
The Red Hills salamander is burrowing species, and is seldom seen out of its underground refuge. Their borrows often open in leaf-litter-free areas near the base of a tree or under siltstone outcroppings. These burrows re often constructed in the lower two-thirds of a steep slope, with the average slope gradient being 50?. Burrows are subject to constant erosion and burrow openings may only last a few months, at which point the salamander must repair or reconstruct its burrow. This species only emerges from its burrow at night, but retreats quickly if disturbed. These salamanders are often observed sitting at the entrance of their burrows with just their head visible on humid nights. They probably forage opportunistically, feeding on any prey item that passes their burrow’s entrance or enters the burrow. The diet of the Red Hills salamander is composed of a variety of invertebrates, including snails, beetles, ants, millipedes and spiders. It is not known whether this species defends its burrows, and sightings of different Red Hills salamanders within a single burrow suggests otherwise. Males may defend territories during the breeding season, but this has not been confirmed.

Egg clusters of this species have not been found in nature, but egg deposition presumably takes place within the burrow system or in deep interconnecting rock fissures underground. The embryos develop directly within the egg capsule and do not pass through a free-living larval phase, hatching out as miniature adults. The Red Hills salamander is an entirely terrestrial (ground-dwelling) species and is not dependent upon water bodies for its reproduction. This species is not water-dwelling at any stage during its life. Courtship, mating, and egg deposition are all thought to occur within the burrow systems. There is no information available on courtship behavior, but fertilization is known to be internal, as makes deposit a packet of sperm called a “spermatophore” which is picked up by the female’s cloaca (urino-genital reproductive opening) so that her eggs may be fertilised within the body cavity. Egg laying is assumed to commence around April, although the breeding season may extend for several months, possibly until September. Clutch size estimates range from 4-16 eggs and parental care is unknown, although many desmognathine salamanders remain with their eggs during development. In the single case of a Red Hills salamander lay eggs in captivity, the female did not brood her eggs and none of them developed.

The Red Hills salamander may have a longevity of up to 11 years, and studies have suggested that females may take 6 years to reach reproductive maturity. These salamanders are active all year round, but they probably retreat deep into the burrow system during periods of extreme cold or drought. The period of greatest activity is spring and summer. Direct predation of this species has not been observed, but small mammals (such as shrews) and reptiles likely eat this species. Feral pigs and armadillos are common in the Red Hills and do considerable damage to the steep ravines inhabited by Red Hills salamanders. They undoubtedly dig out and eat salamanders whenever possible. The burrow is the primary defense of the Red Hills salamander, which is not known to possess poisonous skin glands or exhibit defensive postures. However, this species is capable of biting and will spin when grabbed in an attempt to free itself.
The Red Hills salamander is found on the slopes of damp, shaded ravines with a forest canopy consisting of broad-leaved deciduous trees that provide shade and retain high humidity. The vegetation is dominated by hardwood forest, comprising woodland species such as the big-leaf magnolia, southern magnolia, mountain laurel and oak-leaf hydrangea. This species is often found burrowing in moderately steep areas with a northern exposure, and most often on high, steep, uncut slopes with a high soil moisture content and full tree canopy.

Its ravine habitats are located in the Tallahatta and Hatchetigbee geological formations between the Alabama and Conecuh Rivers in the Red Hills region of southern Alabama, U.S.A. Males and females occupy similar habitats and are seldom seen outside of their burrows. The topsoil of typical habitat is a soft, sandy loam which permits burrowing. This species can tolerate selective logging as long as the ground is not roller-chopped or otherwise prepared, disturbing the burrows of this species.
This species can be found in the Red Hills of south-central Alabama, United States, between the Alabama and Conecuh rivers. It is restricted to the Tallahatta and Hatchetigbee geological formations. The total range area is about 964 km sq., although populations of the Red Hills salamander are patchily distributed depending on appropriate habitat conditions. They have been found in five counties of south-central Alabama: Butler, Conecuh, Covington, Crenshaw, and Monroe.
Population Estimate
Population estimates are difficult to generate due to the secretive, burrowing habits of this species and the isolated location of its populations. It is not rare in good habitat, but current evidence indicates significant losses over the estimated 255 km sq. of formerly occupied habitat. About 60 percent of the occupied habitats is currently owned or leased by paper companies which primarily use a clear-cut system of forest management. This technique, coupled with mechanical site preparation for replanting, appears to completely destroy the habitat for the Red Hills Salamander, and the recovery of affected areas is slow.
Population Trend
This species is thought to be in decline by the ICUN Red List of Threatened Species.
The Red Hills salamander is listed as Endangered in the IUCN Red List of Threatened Species because its extent of occurrence is less than 5,000 km sq. and its area of occupancy is less than 500 km sq., its distribution is severely fragmented, and there is a continuing decline in the extent and quality of its forest habitat in Alabama. Because the Red Hills Salamander lives in a very small section of Alabama, is experiencing habitat loss and has a low reproductive rate, it was placed under Federal protection in 1976. Under the Endangered Species Act, the United States Fish and Wildlife Service listed the Red Hills Salamander as "Threatened".
The habitat of this species has been reduced by timber harvest, the conversion of damp ravines into pine plantations, and the clearance of ridge tops above ravines. All of these practices have destroyed and/or degraded Red Hills salamander habitat. Over collecting may have caused a decline in some areas, especially for scientific purposes and museum specimens up to the 1970s. Around 60% of suitable habitat is on private timber company lands, and detrimental forestry practices continue that disturb the soil and therefore the burrows of Red Hills salamanders. However, some problems have been alleviated by management agreements. Lastly, feral pigs are a threat in localised areas because they dig up salamanders and eat them.
Conservation Underway
This species does not occur in any officially protected areas, although 3 areas (totaling less than 6 hectares) have been set aside to support a limited population. Two of these areas are publicly owned: Lookout Hill Fire Tower (Alabama Forestry Commission) and Haines Island (US Army Corps of Engineers). A cooperative agreement to protect critical habitats is in place. This species is protected as “Threatened” under the US Endangered Species Act of 1973, and is listed as a protected non-game species by the State of Alabama.

In addition, six Habitat Conservation Plans (HCPs) for populations of Red Hills salamanders (covering approximately 25,169 hectares) have been approved by the U.S. Fish and Wildlife Service in cooperation with timber companies in southern Alabama. The goals of the Red Hills salamander HCPs are to allow for timber harvesting while promoting species conservation.
Conservation Proposed
The U.S. Fish and Wildlife Service has made a number of recommendations to facilitate the conservation of the Red Hills salamander in Alabama. This include: the avoidance of clear cutting on slopes containing Red Hills salamander burrows, and especially on the steeper slopes; the avoidance of mechanical site preparation that destroys this species’ burrows and desiccates the soil; the maintenance of woody litter to provide shade and maintain soil moisture and invertebrate content of the soil; the recommendation that two-thirds shade cover be maintained in areas that are selectively cut; the maintenance of a habitat buffer on slopes containing Red Hills salamander burrows in areas where tree felling is occurring; and lastly the avoidance of any chemical sprays within the habitat of this species.

Researchers (e.g. C.K. Dodd) have also made a series of suggestions that would benefit the conservation of the Red Hills salamander. Many of these seem to have been adopted by the U.S. Fish and Wildlife Service in their recommendations, although additional ideas include: allowing hardwoods to regenerate on previously cut and selectively cut slopes containing salamander burrows; protecting the species from collection; and the periodic assessment of Red Hills salamander habitat, especially to ensure that HCP provisions are being honoured and that salamander habitat is protected during forestry operations.
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