The green salamander (Aneides aeneus) is a species of lungless salamander in the family Plethodontidae. It and the Hickory Nut Gorge green salamander (A. caryaensis) are the only currently-described members of the genus Aneides that inhabit any areas in the eastern half of United States (all other Aneides salamanders are found west of the Mississippi River). Rarely seen in the field, the green salamander is an extremely habitat-specific species that is seldom found away from its preferred surroundings: moist, shaded rock crevices.
Description and taxonomyEdit
The green salamander is small and notably flat. Aneides aeneus‘ green, lichen-like blotches against a darker dorsum make it the only salamander in North America with green markings. A. aeneus possesses squared toe-tips, large, conspicuous eyes, and a light blue to yellow ventral surface. Adults range from 8–12 cm (3 to 5 in) with 14 to 15 costal grooves.
Aeneus was formerly considered the only species of the Aneides genus found in the Eastern United States. However, some claimed that there could be up to four different species of Aneides between the Cumberland Plateau and Blue Ridge Escarpment populations. A 2019 study found several A. aeneus populations to represent distinct taxonomic groups, supporting the presence of a possible species complex. At least one was found to represent a distinct species in its own right, the Hickory Nut Gorge green salamander (A. caryaensis). The subgenus Castaneides was created to contain all eastern species of Aneides, with caryaensis being the most basal member of the complex. Castaneides diverged from the Aneides hardii lineage between 27.2 and 32.3 million years ago.
Aneides aeneus is known to inhabit both the Alleghenies and Cumberland Plateau, reaching from southwestern Pennsylvania to northern Alabama and northeastern Mississippi. It is also commonly found in South-Central Ohio. Isolated populations are known at the Blue Ridge Escarpment at the junction of Georgia, North Carolina and South Carolina. The range of the Alleghenies and Cumberland Plateau extends southwest from Fayette County, Pennsylvania through eastern West Virginia, eastern Kentucky, the extreme western portions of Virginia, Tennessee, and Alabama. The population discovered in 1930 in the Great Smoky Mountains National Park has not been located since. There may be disjunct populations on Clinch Mountain, on Bays Mountain and the Appalachian Ridge and Valley, and in the Inner Central Basin of Tennessee (Redmond and Scott, 1996).
Aneides aeneus is an extreme habitat specialist. This species of Aneides is found almost exclusively in an environment following these guild-lines: A. aeneus prefers the crevices of rocks on the sides of cliffs or other outcroppings. Most crevices in which A. aeneus is observed, there is little to no sun light allowed in. This shade may be due to either vegetation outside of the crevice or outcroppings above the crevice. While a number of instances of Aneides aeneus observations in arboreal situations have occurred, most of these instances are in trees either adjacent to rock outcroppings and cliffs, or the specimen in question was located underneath the bark of the tree.
Males are extremely territorial toward other salamanders and would-be predators when disturbed or presented with any manner of threat. Cupp observed aggressive behavior in 45 of 49 instances where a male was placed within an artificial territory of another. Such a high level of aggression is rare in salamanders, and is observed in few other species, though hardly to the degree as observed in A. aeneus. This aggressive behavior, although in different forms, can also be observed in brooding of the female A. aeneus over her eggs. While the male A. aeneus will attack would-be invaders with such actions as butting, snapping, biting or snout-pressing, females will often snap at objects placed within the breeding crevice or near the eggs she guards.
- Aneides aeneus has been observed to begin its period of mating and courtship in late May to early June. For Aneides aeneus, the male will enter the breeding crevices and await the arrival of females. Once a male and female meet, the courtship begins. Like some salamanders of genus Plethodon, A. aeneus begins courtship with the female straddling the base of the male's tail. In this position, and with periodic nudging and encouragement from the male, the two will traverse a small circle within the rock crevice. After some time in this dance, the male will deposit a spermatophore upon the ground and the female, still straddling the base of the male, eventually makes her way to it and “scoops” it up by lateral undulations and slow movements of the base of the tail of the female.
- Eggs are laid soon after courtship. The female A. aeneus lies on her back within the rock crevice, her ventral surface pressed against the ceiling. Egg-laying often takes a period of 20 to 30 hours, where the female will apply an adhesive substance to the surface of the rock, followed by a single egg. Clutches of eggs average 15-25 eggs. The female will stay with the eggs, usually wrapping her body around the cluster or at least pressed against it, guarding them for the entirety of development. When presented with a foreign object, be it a wire, stick, or a wandering insect, the female will attack, eating the invader if possible. Female A. aeneus are not known to feed during brooding. Individuals taken from their eggs had their stomachs and small intestines examined and were all found to be completely empty in Gordon's study in 1971. This guarding period usually lasts 3 months, where the eggs hatch in September.
Dispersal and aggregationEdit
- Eggs of Aneides aeneus hatch throughout the month of September. Aneides aeneus is a direct developing salamander, which means it does not have a larval stage and develops to its adult phase within the egg. Juvenile A. aeneus emerge from the eggs resembling their parents, and will likely leave the crevice in which they were born within 2 months. Following the hatching of their young, the female A. aeneus no longer shows the hyper-aggressiveness expressed during guarding. In fact, it has been observed that the females will do little to stop the collection or otherwise disturbance of her brood after they have hatched from the eggs. Newborn A. aeneus almost always leave the crevice in which they were born and do not return.
- A period in the annual cycle of Aneides aeneus called the pre-hibernation aggregation takes place in which A. aeneus will disperse from the breeding crevices. It is during this period, throughout the month of November, that most specimens of A. aeneus can be observed wandering over and between rock crevices. Following this dispersal, it is thought that A. aeneus ventures deep within the interconnected crevices of the cliffs and rock outcroppings to hibernate. Attempts were made to find A. aeneus underground, beneath logs, within rotten logs, under rocks and under tree bark, but not a single specimen was produced.
Post-hibernation aggregation and dispersalEdit
- Aneides aeneus emerges from hibernation around the month of May. During this period, A. aeneus is observed wandering about the rock crevices and outcroppings, often during light rains at night.
Efforts aimed towards the conservation of such a secretive organism are proving complicated. While the fact that Aneides aeneus is such a habitat-specific salamander results in more vulnerability to habitat destruction, the cliffs and outcroppings it has chosen are relatively safe from harm. It has been speculated that A. aeneus inhabited the ancient chestnut forest that covered a large percentage of the Great Smoky Mountains National Park. It is not known whether or not A. aeneus utilized these trees more or less than its currently preferred habitat, but it is certainly a possibility. In Indiana, the green salamander is listed as an endangered species.
- Geoffrey Hammerson (2004). "Aneides aeneus". IUCN Red List of Threatened Species. 2004: e.T1282A3385547. doi:10.2305/IUCN.UK.2004.RLTS.T1282A3385547.en.
- Frost, Darrel R. (2013). "Aneides aeneus (Cope and Packard, 1881)". Amphibian Species of the World 5.6, an Online Reference. American Museum of Natural History. Retrieved 26 October 2013. CS1 maint: discouraged parameter (link)
- Conant, Roger et al. A Field Guide to Reptiles and Amphibians of Eastern and Central North America. Third Edition, Expanded. Houghton Mifflin Company, Singapore. 1998.
- Gordon, Robert E. A Contribution to the Life History and Ecology of the Plethodontid Salamander Aneides aeneus. American Midland Naturalist. Vol. 47, No. 3 (May 1952), pp. 666-701.
- Dobb, Kenneth C. The Amphibians of Great Smoky Mountains National Park. University of Tennessee Press, Knoxville, TN. 2004.
- Patton, Austin; Apodaca, Joseph J.; Corser, Jeffrey D.; Wilson, Christopher R.; Williams, Lori A.; Cameron, Alan D.; Wake, David B. (December 2019). "A New Green Salamander in the Southern Appalachians: Evolutionary History of Aneides aeneus and Implications for Management and Conservation with the Description of a Cryptic Microendemic Species". Copeia. 107 (4): 748–763. doi:10.1643/CH-18-052. ISSN 0045-8511.
- Snyder, D.H. The Green Salamander Aneides aeneus in Tennessee and Kentucky, With Comments on the Carolina's Blue Ridge Populations. Journal of the Tennessee Academy of Science 66, 1991, pp. 165–169.
- Gordon, Robert E. and Smith, Richard L. Notes on the Life History of the Salamander Aneides aeneus. Copeia. American Society of Ichthyologists and Herpetologists Vol. 1949, No. 3 (September 15, 1949), pp. 173-175.
- Cupp, Paul V. Jr. Territoriality in the Green Salamander, Aneides aeneus. Copeia. American Society of Ichthyologists and Herpetologists. Vol. 1980, No. 3 (September 6, 1980), pp. 463-468.
- Cupp, Paul V. Jr. Fall Courtship of the Green Salamander, Aneides aeneus Herpetologica. Herpetologists' League. Vol. 27, No. 3 (September 1971), pp. 308-310.
- Indiana Legislative Services Agency (2011), "312 IAC 9-5-4: Endangered species of reptiles and amphibians", Indiana Administrative Code, retrieved 28 April 2012 CS1 maint: discouraged parameter (link)