Northern Dusky Salamander
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''Desmognathus fuscus'' is a species of amphibian in the family Plethodontidae ( lungless salamanders). The species is commonly called the dusky salamander or northern dusky salamander to distinguish it from populations in the southern United States which form several distinct species, the southern dusky salamanders ('' D. auriculatus, D. valentinei''). The northern dusky salamander is the most widespread representative of its genus in Canada. It can be found in eastern
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
from extreme eastern
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...
in
New Brunswick New Brunswick (french: Nouveau-Brunswick, , locally ) is one of the thirteen provinces and territories of Canada. It is one of the three Maritime provinces and one of the four Atlantic provinces. It is the only province with both English and ...
south to
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 size of the species' total population is unknown, but is assumed to easily exceed 100,000. The species' habitat differs somewhat geographically; dusky salamanders in the northern part of the range prefer rocky woodland streams, seepages, and springs, while those in the south favor floodplains,
sloughs A slough ( or ) is a wetland, usually a swamp or shallow lake, often a backwater to a larger body of water. Water tends to be stagnant or may flow slowly on a seasonal basis. In North America, "slough" may refer to a side-channel from or feedi ...
, and muddy places along upland streams. They are most common where water is running or trickling. They hide under various objects, such as leaves or rocks, either in or near water. Alternatively, they may enter burrows for protection. The dusky salamander lays its eggs close to water under moss or rocks, in logs, or in stream-bank cavities. The
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
l stage which follows is normally aquatic.


Taxonomy

The spotted dusky salamander (''D. conanti'') and the
flat-headed salamander The flat-headed salamander (''Desmognathus planiceps'') is a species of salamander in the family Plethodontidae. It is endemic to the state of Virginia in the United States. Taxonomy ''Desmognathus planiceps'' was described in 1955 but was lat ...
(''D. planiceps'') were described in the 1950s but were later thought to be synonymous with the northern dusky salamander, but further studies have found all of them to be distinct species from one another.


Description

A small but sturdy
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 ten ...
, the upper body of the northern dusky salamander varies in colour from reddish-brown to gray or olive, with a white or grey underside. The body is sparsely covered with dark spots or
mottling Mottle is a pattern of irregular marks, spots, streaks, blotches or patches of different shades or colours. It is commonly used to describe the surface of plants or the skin of animals. In plants, mottling usually consists of yellowish spots o ...
concentrated on the sides. It also has a light
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal co ...
stripe or two dark stripes that continue on to the first part of the tail. Juvenile colouring consists of five to eight pairs of dorsal spots or blotches located between the front and hind legs. Older individuals tend to be uniformly dark brown or black. As with all dusky salamander species, both juveniles and adults have a pale single stripe outlined in black that extends from the eye and runs diagonally to the rear of the jaw. Additionally, both have 14 costal grooves, larger hind limbs than
forelimb A forelimb or front limb is one of the paired articulated appendages (limbs) attached on the cranial ( anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used instead. ...
s, and a keeled (knife-like) tail that is triangular in cross-section and compressed laterally at the base. The tail is less than half its body length and is normally lighter in colour in comparison to the body. Adults attain lengths of up to 14 cm, with the average length of adult males and females being 9.4 cm and 8.6 cm, respectively. Being from the family Plethodontidae, the northern dusky salamander is lungless. It absorbs oxygen through the skin and membranous tissue located in the mouth and throat. The dusky salamander also has a naso-labial groove, which aids
olfaction The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...
, and thus the ability to search out mates and prey through smell.


Similar species

The dusky salamander is similar in appearance to and thus often confused with the
Allegheny Mountain dusky salamander The Allegheny Mountain dusky salamander (''Desmognathus ochrophaeus'') is a species in the Plethodontidae ( lungless salamander) family. It is found in the eastern United States, in the Niagara Glen Nature Reserve adjacent to the lower Niagara R ...
(''Desmognathus ochrophaeus''). Distinguishing characteristics are that the dorsal spots of the Mountain dusky salamander are usually
chevron Chevron (often relating to V-shaped patterns) may refer to: Science and technology * Chevron (aerospace), sawtooth patterns on some jet engines * Chevron (anatomy), a bone * '' Eulithis testata'', a moth * Chevron (geology), a fold in rock ...
-shaped and its tail rounded at the base rather than laterally compressed. Additionally, hybridization has been known to occur between the Allegheny Mountain dusky salamander and the northern dusky salamander. The northern dusky salamander can also be differentiated from other lungless salamanders including the eastern red-backed, the northern two-lined and the four-toed. They have a distinguishing pale-coloured line that runs from behind their eyes to the rear of the jaw, and heavier set bodies with longer hind legs than front legs.


Range

This species is native to North America, and occurs throughout central-eastern regions of Canada and the United States, from southern
New Brunswick New Brunswick (french: Nouveau-Brunswick, , locally ) is one of the thirteen provinces and territories of Canada. It is one of the three Maritime provinces and one of the four Atlantic provinces. It is the only province with both English and ...
, southeastern
Quebec Quebec ( ; )According to the Canadian government, ''Québec'' (with the acute accent) is the official name in Canadian French and ''Quebec'' (without the accent) is the province's official name in Canadian English is one of the thirtee ...
and southern
Ontario Ontario ( ; ) is one of the thirteen provinces and territories of Canada.Ontario is located in the geographic eastern half of Canada, but it has historically and politically been considered to be part of Central Canada. Located in Central Ca ...
south to
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 = ...
. Populations south of this range are now thought to belong to other, previously-synonymized species such as '' D. conanti''. The Canadian distribution accounts for approximately 5% of the global range. Within its Canadian range, the northern dusky salamander usually occurs in forested
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 ...
located in high elevation, low-order streams. There are two separate units (DU), the Quebec/New Brunswick DU and the Carolinian DU in Ontario. Ohio populations are reviewed in "Amphibians of Ohio" (2013). An isolated breeding population exists in eastern Michigan; however, it is not known whether this population is natural or was introduced.


Habitat

In the northern extent of their range, the northern dusky salamander inhabits saturated soil near springs, seepages, and small tributaries of small headwater streams otherwise known as the
riparian zone A riparian zone or riparian area is the interface between land and a river or stream. Riparian is also the proper nomenclature for one of the terrestrial biomes of the Earth. Plant habitats and communities along the river margins and banks ar ...
.
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 ...
quality is optimal in undisturbed watersheds and where water is running or trickling and there is an abundance of forest cover The forest cover serves to keep the water cool and well oxygenated, and maintains moisture and temperature at levels necessary for salamander survival. In the south, the northern dusky salamander can be found in upland streams as well as floodplains, sloughs and muddy sites.


Behavior and ecology

The species uses subterranean retreats or
burrow An Eastern chipmunk at the entrance of its burrow A burrow is a hole or tunnel excavated into the ground by an animal to construct a space suitable for habitation or temporary refuge, or as a byproduct of locomotion. Burrows provide a form of sh ...
s near the streams edge as well as leaf litter, logs, rocks and moss as a source of protective cover for avoiding
desiccation Desiccation () is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. ...
and
predators Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
. These microhabitats are also important for foraging and nesting both of which take place on land close to the water's edge. As a result of desiccation and predators, activity of the northern dusky salamander peaks in the morning, and the evening and early night. Alongside the stream, females nest in cryptic microhabitats where soil is saturated with water. Their eggs are often laid in logs, under rocks or moss, or stream-bank cavities. During development while in the
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
l stage, the northern dusky salamander is strictly aquatic, its habitat the interstitial spaces between rocks of the streambed. In winter, they remain in shallow running water, whereas adults overwinter in subterranean retreats or in streams, often remaining active throughout winter if the substrate doesn't freeze. As a relatively small amphibian, the northern dusky salamander spends most of its life in hiding. If predated it is capable of
autotomy Autotomy (from the Greek language, Greek ''auto-'', "self-" and ''tome'', "severing", wikt:αὐτοτομία, αὐτοτομία) or self-amputation, is the behaviour whereby an animal sheds or discards one or more of its own appendages, usual ...
along any point of its tail, but lacks chemical defense mechanisms against its main predators which include larger salamanders, birds, fish, snakes, crayfish, and small mammals. known as maybe rodents or mice The home range of the northern dusky salamander is limited to 1m2 to 3.6 m2. The northern dusky salamander has seasonal variations with its patterns of movement. During warmer months the salamanders have larger home ranges that average around 1.5 m2. This is reduced during the winter and some populations move into specific areas for condensed winter retreats.


Reproduction and development

Due to their lack of mobility, some populations of dusky salamander are genetically distinct. Breeding is terrestrial and occurs annually in spring or fall and includes elaborate courtship rituals. Adult males have papillose
cloaca In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles and birds, a ...
l lips and a small mental gland. These glandular tissues become enlarged when sexually active. Female individuals lack a mental gland and have folded cloacal lips. For reproduction, the male applies the snout, cheeks and mental gland to the snout of the female, who usually responds by picking up the
spermatophore A spermatophore or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especially salamanders and arthropods, and transferred in entirety to the female's ovipore during reproduction. Spermatophores ...
.
Fecundity Fecundity is defined in two ways; in human demography, it is the potential for reproduction of a recorded population as opposed to a sole organism, while in population biology, it is considered similar to fertility, the natural capability to pr ...
increases with body size. Females normally deposit between 10 and 30 eggs under logs, moss or rocks located streamside where soil is saturated with water.
Clutch size __NOTOC__ A clutch of egg (biology), eggs is the group of eggs produced by birds, amphibians, or reptiles, often at a single time, particularly those laid in a nest. In birds, destruction of a clutch by predators (or removal by humans, for exam ...
has been known to vary geographically and can be as large as forty-five, or as few as eight. Females remain with their eggs for an incubation period of six to ten weeks (45 to 60 days) in order to protect them from desiccation and predation . The larvae are predominantly aquatic and approximately 1.5 cm in length upon hatching. The larvae then
metamorphose Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some insec ...
into semi-terrestrial adults, with juvenile salamanders being 2.8 to 4.4 cm in length. Their
biphasic Biphasic, meaning having two phases, may refer to: * Phase (matter) In the physical sciences, a phase is a region of space (a thermodynamic system), throughout which all physical properties of a material are essentially uniform. Examples ...
life cycle includes an aquatic state of seven to 16 months, followed by a semi-terrestrial adult stage. The northern dusky salamander attains
sexual maturity Sexual maturity is the capability of an organism to reproduce. In humans it might be considered synonymous with adulthood, but here puberty is the name for the process of biological sexual maturation, while adulthood is based on cultural definitio ...
at approximately three to four years of age.
Life expectancy Life expectancy is a statistical measure of the average time an organism is expected to live, based on the year of its birth, current age, and other demographic factors like sex. The most commonly used measure is life expectancy at birth ...
is 10 to 15 years.


Feeding

The northern dusky salamander is considered to be feeding generalist, with its diet based on food availability. Stream salamanders are known to be significant predators. This is namely because they are dominant vertebrates within headwater riparian forest ecosystems, with a biomass greater than that reported for fish, birds or small mammals. The species is carnivorous and consumes a variety of aquatic and terrestrial invertebrates. Larvae feed predominantly on aquatic invertebrates (such as copepods, other crustaceans and insect larvae), as well as mites, whereas the adult diet consists of 60 to 85% of terrestrial invertebrates, including arthropods (such as crustaceans, spiders, mites, flies and fly larvae, ants, beetles and beetle larvae, centipedes, moths and mayflies), snails, slugs and earthworms. When prey is in excess, the northern dusky salamander does typically have a preference for the larger and fleshier terrestrial invertebrates, such as earthworms. The aquatic portion of the adult's diet is habitat specific and commensurate with the seasonal abundance and diversity of invertebrates.


Conservation


Population and trends

Current data does not allow an accurate estimate of
population size In population genetics and population ecology, population size (usually denoted ''N'') is the number of individual organisms in a population. Population size is directly associated with amount of genetic drift, and is the underlying cause of effect ...
or trends. That said, the total adult population size of the northern dusky salamander is known to exceed 100,000 individuals. There are numerous stable populations throughout the range. The species is widespread in Quebec and New Brunswick but local densities are usually low. In Ontario, the species is rare with a population size estimated at fewer than 250 individuals.


Threats and limiting factors

The northern dusky salamander is extremely vulnerable to
desiccation Desiccation () is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. ...
and therefore reliant on clean headwater streams. Resultantly, contamination of ground water or waterways through pollution from
urban area An urban area, built-up area or urban agglomeration is a human settlement with a high population density and infrastructure of built environment. Urban areas are created through urbanization and are categorized by urban morphology as cities, t ...
s, industry, or
agriculture Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people to ...
, can be catastrophic to local populations. The disappearance of the species from the
Acadia National Park Acadia National Park is an American national park located along the mid-section of the Maine coast, southwest of Bar Harbor. The park preserves about half of Mount Desert Island, part of the Isle au Haut, the tip of the Schoodic Peninsula, and ...
in Maine is believed to be the result of heavy metal contamination. Freshwater stream
acidification Acidification may refer to: * Ocean acidification, decrease in the pH of the Earth's oceans * Freshwater acidification, atmospheric depositions and soil leaching of SOx and NOx * Soil acidification, buildup of hydrogen cations, which reduces the ...
also poses a significant threat with 40% of streams in the southern
Appalachians The Appalachian Mountains, often called the Appalachians, (french: Appalaches), are a system of mountains in eastern to northeastern North America. The Appalachians first formed roughly 480 million years ago during the Ordovician Period. They ...
showing signs of acidification. Changes to stream flow or the groundwater supply, can have significant impacts on local salamander genetics and populations vis-à-vis loss of suitable aquatic or terrestrial habitat, bank instability from excessive
runoff Runoff, run-off or RUNOFF may refer to: * RUNOFF, the first computer text-formatting program * Runoff or run-off, another name for bleed, printing that lies beyond the edges to which a printed sheet is trimmed * Runoff or run-off, a stock market ...
, or simply changes to the moisture in the terrestrial habitat. Such changes can be naturally occurring or artificially induced (e.g. discharged water volumes). The activities of
forestry Forestry is the science and craft of creating, managing, planting, using, conserving and repairing forests, woodlands, and associated resources for human and environmental benefits. Forestry is practiced in plantations and natural stands. Th ...
can be similarly devastating. Timber harvesting,
wind farm A wind farm or wind park, also called a wind power station or wind power plant, is a group of wind turbines in the same location used Wind power, to produce electricity. Wind farms vary in size from a small number of turbines to several hundre ...
s and watershed urbanization reduce water supply, water quality and microhabitat availability. Aquatic habitat can be degraded through
siltation Siltation, is water pollution caused by particulate terrestrial clastic material, with a particle size dominated by silt or clay. It refers both to the increased concentration of suspended sediments and to the increased accumulation (temporary or ...
of streams, or the microhabitat conditions of the forest floor undergoing alterations. Siltation is of particular consequence to the northern dusky salamander because the interstitial spaces that they use for foraging, nesting and
overwintering Overwintering is the process by which some organisms pass through or wait out the winter season, or pass through that period of the year when "winter" conditions (cold or sub-zero temperatures, ice, snow, limited food supplies) make normal activi ...
are lost. Urbanization has resulted in the disappearance of the species in Mount Saint-Hellaire National Park in Quebec, as well as other areas. Vulnerability to
extirpation Local extinction, also known as extirpation, refers to a species (or other taxon) of plant or animal that ceases to exist in a chosen geographic area of study, though it still exists elsewhere. Local extinctions are contrasted with global extinct ...
is further heightened when the species relies on a single watershed. Such is the case of the Ontario population of the northern dusky salamander. The species is also threatened through the introduction of predatory fish, such as
Brook Trout The brook trout (''Salvelinus fontinalis'') is a species of freshwater fish in the char genus ''Salvelinus'' of the salmon family Salmonidae. It is native to Eastern North America in the United States and Canada, but has been introduced elsewhere ...
.


Conservation status

The northern dusky salamander is listed as endangered in Ontario and is declining in many parts of the United States yet some populations remain stable. The International Union for Conservation of Nature (IUCN) lists its global status as Least Concern. In the province of Ontario, where the status of the Carolinian population of the northern dusky salamander is listed as Endangered, the northern dusky salamander is protected under the Ontario Endangered Species Act of 2007 and by the Ontario Fish and Wildlife Conservation Act. These acts protect its habitat and make it illegal to possess, harm or kill the species. In Ontario, a dusky salamander recovery team entitled the "Allegheny Mountain Dusky Salamander and Northern Dusky Salamander Recovery Strategy" has also been established to develop a recovery plan for both species. The Quebec / New Brunswick population of the northern dusky salamander is considered not at risk. Nonetheless, it is currently on a list of species likely to be designated as
threatened Threatened species are any species (including animals, plants and fungi) which are vulnerable to endangerment in the near future. Species that are threatened are sometimes characterised by the population dynamics measure of ''critical depensat ...
or vulnerable by the Quebec provincial government and is also protected by a provincial act that prohibits the collecting or selling of specimens. Likewise, Article 22 of the provincial Environmental Quality Act offers protection against unregulated degradation of the dusky salamander's environment. In New Brunswick, the species is designated as Sensitive under the General Status of Species in Canada. Protection is offered the species by the New Brunswick Fish and Wildlife Act.


References


External links


Dusky Salamander (''Demognathus fuscus'')
Natural Resources Canada {{Taxonbar, from=Q733067 fuscus Amphibians of Canada Amphibians of the United States Fauna of the Great Lakes region (North America) Fauna of the Northeastern United States Fauna of the Southeastern United States Ecology of the Appalachian Mountains Cenozoic amphibians of North America Extant Pleistocene first appearances Pleistocene animals of North America Pleistocene United States Amphibians described in 1820 Taxa named by Constantine Samuel Rafinesque Taxonomy articles created by Polbot