Red Hills Salamander

Phaeognathus hubrichti is an elongated salamander with reduced limbs and a prehensile tail. P. hubrichti is uniform dark brown in coloration and lacks the light line from the eye to the jaw of other desmognathine salamanders. Elongation in P. hubrichti is due to an increased number of trunk vertebrae, 22 vertebrae in P. hubrichti while most other Desmognathines have 15 (Highton 1961).At one time, Phaeognathus hubrichti was considered possibly threatened by overcollection for museum specimens and subsequent destruction of habitat (Mount and Schwaner 1970). After its initial discovery under leaf litter at the type locality, herpetologists were unable to find additional specimens for some time. Valentine (1963a) suggested a collecting technique of locating salamanders in their burrow entrances and then driving a pick behind the salamander to prevent its escape down the burrow. Although obviously destructive to the fragile ravine habitat, this technique produced many museum specimens. Mount and Schwaner (1970) expressed concern at the destruction of the habitat of the type locality by collectors and suggested a new technique of "fishing" for P. hubrichti at the entrance to their burrows. A small fishhook baited with a cricket or spider is dangled in front of a P. hubrichti burrow and once the bait is taken the salamander can be gently pulled from its burrow. This technique rarely causes harm to salamanders and has been successfully employed in other studies (Gunzburger 1999). Due to its listing as a federally threatened species, collection of P. hubrichti is now prohibited.

Phaeognathus hubrichti is restricted steep mesic ravine habitats in the Tallahatta and Hatchetigbee geological formations between the Alabama and Conecuh Rivers in the Red Hills physiographic region of southern Alabama, USA (Schwaner and Mount 1970). Populations of P. hubrichti are patchy depending on appropriate habitat conditions. P. hubrichti have only been found in five counties: Butler, Conecuh, Covington, Crenshaw, Monroe. P. hubrichti was described by Highton (1961) based on the collection of the holotype by Dr. Leslie Hubricht and then redescribed by Valentine (1963a). Morphological features such as elongate form, reduced limbs, and eyelid morphology suggested Phaeognathus hubrichti was a burrowing salamander (Valentine 1963a). Habitat preferred by Phaeognathus hubrichti is steep, mesic ravines shaded by a mature hardwood overstory (Jordan 1975; Schwaner and Mount 1970) P. hubrichti is almost entirely fossorial and appears to be found most often in areas of soft soil which allows burrowing. P. hubrichti burrows are often most abundant on the steepest regions of slopes (Valentine 1963b). Burrows are subject to constant erosion and burrow openings may only last a few motnhs, at which point the salamander must repair or reconstruct its burrow (Gunzburger and Guyer 1998).

Concern about the impact of forestry operations on the long-term survival of Phaeognathus hubrichti populations was first suggested by Jordan and Mount (1975). Clearcutting removal of the overstory of ravine habitats totally eliminated P. hubrichti populations (Schwaner and Mount 1978). P. hubrichti was listed by the USFWS as threatened in 1976 due to habitat loss and degradation, mostly through forestry operations (Schwaner and Mount 1978). Dodd (1991) resurveyed sites studied by Schwaner and Mount (1978),indicating that although some P. hubrichti populations appeared stable, many were declining apparently due to continued forestry practices. Dodd (1991) suggested 100-200 m buffer zones be left uncut near ravine slopes to reduce moisture loss.

Because Phaeognathus hubrichti is a fossorial species, little is known of its behaviors and habits. Courtship, mating, and egg deposition probably all occur within burrow systems. There is no information available on courtship behavior, but fertilization is internal by means of a spermatophore. No aquatic larval stage is present. Eggs have never been found in nature, but based on enlarged ova in collected specimens Brandon (1965) suggested females lay eggs in April. Schwaner and Mount (1970) indicate the breeding season may extend for several months as a recently spent female was collected in September. Clutch size estimates from ova of females range from 4-6 (Petranka 1998) to 8-9 (Brandon 1965). A 115 mm SVL captive female laid 16 eggs about 7 mm in diameter (Brandon and Maruska 1982). Skeletochronological age estimates for P. hubrichti indicate individuals may live as long as 11 years and age is positively correlated with size (SVL) (Parham et al. 1996). This study suggested that female P. hubrichti may take 6 years to reach reproductive maturity (Parham et al. 1996). Phaeognathus hubrichti are often observed sitting with just their head visible at the entrance of their burrow on moist nights. P. hubrichti probably forages opportunistically on any prey item that passes by the burrow entrance or enters the burrow. Diet of P. hubrichti consists of a variety of invertebrates including snails, beetles, ants, millipedes, and spiders (Brandon 1965; Gunzburger 1999). Analysis of proteins from individuals from 13 populations of Phaeognathus hubrichti throughout their range suggests that there are 2 distinct geographic groups (McKnight et al. 1991). An effort to conserve population within both of these groups should be attempted.

Because Phaeognathus hubrichti is endemic to Alabama and possesses so many unique and interesting features, it has been named the State Amphibian of Alabama. Thus all citizens of Alabama can be proud of this unique part of their natural heritage and hopefully support its continued protection and conservation. http://archives.state.al.us/emblems/st_amphib.html

Maximum longevity: 19.7 years (captivity)

Red Hills salamanders live for approximately 11 years in the wild, which is determined by counting growth rings on limb bones. They are exceptionally rare and there are no records indicating the average lifespan of captive individuals. However, one female specimen lived for over 6 years in captivity and was able to lay eggs prior to being released.

Average lifespan
Status: captivity: 6 years.

Average lifespan
Status: wild: 11 years.

Primary predators of Red Hills salamanders are thought to include various species of bird, snakes, and mammals such as coyotes and badgers. When threatened by a potential predator, Red Hills salamanders bare their teeth and may attempt to strike. They are also known to "head-butt" burrow intruders. When head-butting, individuals thrust themselves from their burrows with their small limbs, driving their head into the intruder. The intruder is startled upon impact and is knocked out of the entrance and down the hill the burrow was created on. This technique is most effective on intruders that are smaller or of comparable size. If captured, an individual may gyrate in a circular pattern similar to that of a snake, attempting to loosen the attacker's grip. Its primary form of defense, however, is to remain inside its burrow as much as possible.

Known Predators:

• birds, (Aves)
• snakes, (Serpentes)
• mammals, (Mammalia)
• coyotes, (Canis latrans)
• badgers, (Taxidea taxus)

Anti-predator Adaptations: cryptic

Male Red Hills salamanders range from 180 to 220 mm in length, whereas females range in length from 190 to 230 mm. Males can weigh up to 22 g, but most weigh between 7 and 15 g. Females are considerably lighter, weighing between 6 and 12 g, with a maximum of 14 g. Red Hills salamanders belong to the Plethodontinae family, and therefore are lungless and breath through their moist skin. The skin is dark brown in colour with an occasional light spot interspersed around its body, specifically around the facial region. Some larger males have have pale spots on either side of their body, at the base of their tail. Although Red Hills salamanders are significantly longer than their relatives, they have much smaller limbs. The anterior limbs are approximately 11 mm in length and have 4 toes, and the posterior limbs are approximately 14 mm in length and have 5 toes. They have between 20 and 22 costal grooves along the mid-section, which is significantly more than most salamanders. Red Hills salamanders, like other plethodons, have fixed lower jaws, and barely visible nasolabial grooves on their snouts that assist in chemo-reception. Late term pregnant females have eggs that are clearly visible through her skin along each side of her mid-section.

Range mass: 5 to 20 g.

Average mass: 11 g.

Range length: 160 to 250 mm.

Other Physical Features: ectothermic ; heterothermic ; bilateral symmetry

Sexual Dimorphism: sexes shaped differently

Phaeognathus hubrichti prefers the soft-soiled hills of the Red Hills area and seeks out slopes on mesic ravines that are moist, soft and loamy. It also prefers burrow under full canopy of hardwood trees on relatively high and steep ridges sloping down into slow, shallow streams. Most P. hubrichti burrows are found in the center of such slopes, as this location is the least likely to be affected by erosion. It is often found in abandoned burrows of other ground-dwelling species, as it cannot dig effectively itself. If an abandoned burrow is too small, it adjust the size of the burrow by wiggling into the hole and rolling inside the hole until it is large enough for the salamander’s use. Phaeognathus hubrichti burrows are approximately 40 mm wide with a distinctive oval shape. This species is found at an average elevation of 170 m.

Average elevation: 170 m.

Habitat Regions: temperate

Terrestrial Biomes: forest

The geographic range of Red Hills salamanders (Phaeognathus hubrichti) is restricted to the Red Hills region of south central Alabama, USA. They have been documented in only 6 counties in the state of Alabama (Covington, Crenshaw, Monroe, Butler, Barbour, Conecuh). The Alabama River demarcates the species' western-most boundary and the Conecuh River demarcates their eastern-most boundary. Although they appear to reside in adjacent areas to the north and south, their fossorial tendencies make their geographic range difficult to verify. The total area that is occupied by Red Hills salamanders is estimated to be approximately 25,500 hectares.

Biogeographic Regions: nearctic (Native )

Red Hills salamanders are primarily insectivorous, and gut content analyses have revealed a preference for spiders, earthworms, millipedes, beetles, mites and fly larvae. In some instances, they have been seen foraging on snails and molted snake skins. Other foods include fungus and detritus.

Animal Foods: eggs; insects; terrestrial non-insect arthropods; mollusks; terrestrial worms

Other Foods: fungus; detritus

Primary Diet: carnivore (Eats eggs, Insectivore , Eats non-insect arthropods, Molluscivore , Vermivore, Scavenger )

Phaeognathus hubrichti is an important prey item for a number of different vertebrate species, including various species of snakes, mammals, birds, other amphibians. In addition, this species is insectivorous and my help control a variety of insect pest species. There is no information available regarding potential parasites of this species.

Ecosystem Impact: soil aeration

There are no known positive effects of Phaeognathus hubrichti on humans.

Positive Impacts: research and education

There are no known adverse effects of Phaeognathus hubrichti on humans.

Red Hills salamanders sexually mature by 100 mm in length, which is approximately 5 to 6 years of age for females. Males mature by 80 mm in length, which take as little as one year for males. Once eggs hatch, young are similar in appearance to adults, with the exception of their bright red gills, which are laterally positioned between the head and forelimbs. Like their adult counterparts, young also have labial folds, but no eyelids. About 10 days after hatching, young undergo metamorphosis, during which they grow eyelids, lose their labial folds, and reabsorb their gills. Most individuals metamorphose at about 37 mm in length and may have some paleness along the ventral surface after metamorphosis.

Development - Life Cycle: metamorphosis

Red Hills salamanders are classified as "endangered" on the IUCN's Red List of Threatened Species and has been protected by the United States Endangered Species Act as "threatened" since 1976. This species is found only in south central Alabama and is listed as a protected non-game species by the state of Alabama. Greater than 40% of potential habitat is owned or managed by pulp corporations, and although it does not occur in any officially protected habitat, a little more than 6 hectares have been set aside by private and public ownership to support the conservation and management of this species. Unfortunately, evidence suggests that Red Hills salamanders do not readily re-populate reforested areas, making their recovery significantly more difficult. In 2010, the United States Nature Conservancy purchased 723 hectares of the Red Hills Conservation Area in order to support the long-term protection of this species. The Nature Conservancy, the State of Alabama and the United States Fish and Wildlife Service, continue to work together to protect areas of salamander habitat from potential degradation.

US Federal List: threatened

CITES: no special status

State of Michigan List: no special status

IUCN Red List of Threatened Species: endangered

There is little information available regarding communication and perception in Phaeognathus hubrichti. This species likely does not communicate audibly; however, nasolabial grooves on the snout are thought to assist in chemoreception, and the interior groove is lined with glands. Although individuals are thought to aggressively defend burrows, there is no data to confirm this. Numerous male specimens are documented as having bite marks, suggesting aggressive interactions with conspecifics, possibly as a result of competition for mates or burrows.

Communication Channels: visual ; chemical

Perception Channels: visual ; tactile ; chemical

There is no information available regarding the mating system of Phaeognathus hubrichti.

Phaeognathus hubrichti males become reproductively mature in less than a year, whereas females take approximately 6 years to reach reproductive maturity. Phaeognathus hubrichti mates once per year. Once mated, females carry fertilized eggs in their oviducts for approximately 20 days. Eggs are then laid during June or July. After being laid, eggs incubate for another 60 days and then are attached to the walls of the burrow in a space that is moist, but not wet. It typically takes around 20 days for young to hatch and emerge. Females hatch an average of 7 offspring per breeding cycle. Little else is known of the reproductive behavior of this species.

Breeding interval: <<Phaeognathus hubrichti>> mates once per year.

Breeding season: Phaeognathus hubrichti lays its eggs in late June-July.

Average number of offspring: 7.

Average time to hatching: 20 days.

Average age at sexual or reproductive maturity (female): 6 years.

Average age at sexual or reproductive maturity (male): <1 years.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (Internal ); oviparous

Phaeognathus hubrichti mothers stay close to their eggs during development. If the eggs are disturbed, she may ingest them to prevent losing the energy invested in their development. When carrying fertilized eggs, females aggresively defend themselves by biting. Once the eggs are laid, however, she is relatively less aggressive. Paternal care is thought to be non-existent in this species. There is no further information on parental care in P. hubrichti.

Parental Investment: female parental care ; pre-hatching/birth (Protecting: Female)

The Red Hills salamander (Phaeognathus hubrichti) is a fairly large, terrestrial salamander growing to about 255 millimetres (10.0 in). Its body color is gray to brownish without markings, and its limbs are relatively short. It is the official state amphibian of Alabama, the state it is endemic to. It is the only species in the genus Phaeognathus.