Green Salamander
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The green salamander (''Aneides aeneus'') is a species of lungless
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All t ...
in the family
Plethodontidae Plethodontidae, or lungless salamanders, are a family of salamanders. Most species are native to the Western Hemisphere, from British Columbia to Brazil, although a few species are found in Sardinia, Europe south of the Alps, and South Korea. In ...
. It and the
Hickory Nut Gorge green salamander The Hickory Nut Gorge green salamander (''Aneides caryaensis'') is a species of lungless salamander in the family Plethodontidae. It is endemic to the Hickory Nut Gorge in the state of North Carolina in the United States. Taxonomy It is a memb ...
(''A. caryaensis'') are the only currently-described members of the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
''Aneides'' that inhabit any areas in the eastern half of United States (all other ''Aneides'' salamanders are found west of the
Mississippi River The Mississippi River is the second-longest river and chief river of the second-largest drainage system in North America, second only to the Hudson Bay drainage system. From its traditional source of Lake Itasca in northern Minnesota, it fl ...
). Rarely seen in the field, the green salamander is an extremely habitat-specific
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
that is seldom found away from its preferred surroundings: moist, shaded rock crevices.


Description and taxonomy

The green salamander is small and notably flat. ''Aneides aeneus''‘ green, lichen-like blotches against a darker dorsum make it the only
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All t ...
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 Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
found in the Eastern
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territori ...
. 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 In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
. At least one was found to represent a distinct species in its own right, the
Hickory Nut Gorge green salamander The Hickory Nut Gorge green salamander (''Aneides caryaensis'') is a species of lungless salamander in the family Plethodontidae. It is endemic to the Hickory Nut Gorge in the state of North Carolina in the United States. Taxonomy It is a memb ...
(''A. caryaensis''). The subgenus ''
Castaneides Climbing salamanders is the common name for plethodontid (lungless) salamanders of the genus ''Aneides''. It contains 10 species native to North America, distributed between the Pacific Coast (7 species), Sacramento Mountains (1 species), and A ...
'' 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. Members of ''Castaneides'' are the only
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All t ...
s in North America with green markings. Due also to their hyper-specific
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
, they are almost unmistakable when found in the field.


Geographic distribution

''Aneides aeneus'' is known to inhabit both the
Alleghenies The Allegheny Mountain Range (; also spelled Alleghany or Allegany), informally the Alleghenies, is part of the vast Appalachian Mountain Range of the Eastern United States and Canada and posed a significant barrier to land travel in less develo ...
and Cumberland Plateau, reaching from southwestern
Pennsylvania Pennsylvania (; ( Pennsylvania Dutch: )), officially the Commonwealth of Pennsylvania, is a state spanning the Mid-Atlantic, Northeastern, Appalachian, and Great Lakes regions of the United States. It borders Delaware to its southeast, ...
to northern
Alabama (We dare defend our rights) , anthem = "Alabama" , image_map = Alabama in United States.svg , seat = Montgomery , LargestCity = Huntsville , LargestCounty = Baldwin County , LargestMetro = Greater Birmingham , area_total_km2 = 135,765 ...
and northeastern
Mississippi Mississippi () is a state in the Southeastern region of the United States, bordered to the north by Tennessee; to the east by Alabama; to the south by the Gulf of Mexico; to the southwest by Louisiana; and to the northwest by Arkansas. Miss ...
. It is also commonly found in South-Central Ohio. Isolated populations are known at the Blue Ridge Escarpment at the junction of
Georgia Georgia most commonly refers to: * Georgia (country), a country in the Caucasus region of Eurasia * Georgia (U.S. state), a state in the Southeast United States Georgia may also refer to: Places Historical states and entities * Related to the ...
,
North Carolina North Carolina () is a state in the Southeastern region of the United States. The state is the 28th largest and 9th-most populous of the United States. It is bordered by Virginia to the north, the Atlantic Ocean to the east, Georgia and ...
and
South Carolina )'' Animis opibusque parati'' ( for, , Latin, Prepared in mind and resources, links=no) , anthem = " Carolina";" South Carolina On My Mind" , Former = Province of South Carolina , seat = Columbia , LargestCity = Charleston , LargestMetro = ...
. The range of the Alleghenies and Cumberland Plateau extends southwest from
Fayette County, Pennsylvania Fayette County is a county in the Commonwealth of Pennsylvania. It is located in southwestern Pennsylvania, adjacent to Maryland and West Virginia. As of the 2020 census, the population was 128,804. Its county seat is Uniontown. The county wa ...
through eastern
West Virginia West Virginia is a state in the Appalachian, Mid-Atlantic and Southeastern regions of the United States.The Census Bureau and the Association of American Geographers classify West Virginia as part of the Southern United States while the B ...
, eastern
Kentucky Kentucky ( , ), officially the Commonwealth of Kentucky, is a state in the Southeastern region of the United States and one of the states of the Upper South. It borders Illinois, Indiana, and Ohio to the north; West Virginia and Virginia ...
, the extreme western portions of
Virginia Virginia, officially the Commonwealth of Virginia, is a state in the Mid-Atlantic and Southeastern regions of the United States, between the Atlantic Coast and the Appalachian Mountains. The geography and climate of the Commonwealth ar ...
,
Tennessee Tennessee ( , ), officially the State of Tennessee, is a landlocked state in the Southeastern region of the United States. Tennessee is the 36th-largest by area and the 15th-most populous of the 50 states. It is bordered by Kentucky to th ...
, and Alabama. The population discovered in 1930 in the
Great Smoky Mountains National Park Great Smoky Mountains National Park is an American national park in the southeastern United States, with parts in North Carolina and Tennessee. The park straddles the ridgeline of the Great Smoky Mountains, part of the Blue Ridge Mountains, w ...
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).


Life history

''Aneides aeneus'' is an extreme
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
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 Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. The habitats pose nu ...
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 In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
, 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.


Annual cycle


Breeding period

*''Aneides aeneus'' has been observed to begin its period of mating and
courtship Courtship is the period wherein some couples get to know each other prior to a possible marriage. Courtship traditionally may begin after a betrothal and may conclude with the celebration of marriage. A courtship may be an informal and private m ...
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 ''Plethodon'' is a genus of salamanders in the family Plethodontidae. They are also known as woodland salamanders or, more rarely, slimy salamanders. All members of the genus are endemic to North America (Canada and USA). They have no aquatic lar ...
'', ''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 aggregation

*Eggs of ''Aneides aeneus'' hatch throughout the month of September. ''Aneides aeneus'' is a direct developing
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All t ...
, 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.


Hibernation

*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 Hibernation is a state of minimal activity and metabolic depression undergone by some animal species. Hibernation is a seasonal heterothermy characterized by low body-temperature, slow breathing and heart-rate, and low metabolic rate. It most ...
. 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 dispersal

*''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.


Conservation

Efforts aimed towards the conservation of such a secretive
organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and ...
are proving complicated. While the fact that ''Aneides aeneus'' is such a habitat-specific
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All t ...
results in more vulnerability to
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
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 Great Smoky Mountains National Park is an American national park in the southeastern United States, with parts in North Carolina and Tennessee. The park straddles the ridgeline of the Great Smoky Mountains, part of the Blue Ridge Mountains, w ...
. 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. When inhabiting arboreal habitats studies have found that green salamanders prefer hardwood trees to conifers. In
Indiana Indiana () is a U.S. state in the Midwestern United States. It is the 38th-largest by area and the 17th-most populous of the 50 States. Its capital and largest city is Indianapolis. Indiana was admitted to the United States as the 19th s ...
, the green salamander is listed as an endangered species.


References


External links

* * {{Taxonbar, from=Q277791 Aneides Amphibians of the United States Endemic fauna of the United States Amphibians described in 1881