Geographic Range
Populations of
Necturus lewisi
are endemic to North Carolina in the United States. More specifically, these amphibians
reside in parts of Craven, Durham, Franklin, Granville, Johnston, Lenoir, Nash, Orange,
Vance, and Wake counties, around 35°N latitude. They reside in the Neuse and Tar-Pamlico
river basins and research has been conducted on these animals in the Little River,
a tributary of the Neuse River.
Habitat
Necturus lewisi lives in freshwater streams; none have been found in lakes or ponds. Requiring clean, fresh water, populations of this aquatic amphibian are most prevalent in streams that are wider than 15 meters, deeper than 100 cm, and have a flow rate of greater than 10 cm per second. Both juveniles and adults dwell in loose granite rocks and sand or gravel areas as well as under bedrock from the stream banks.
The salamanders develop home retreat areas free of debris and algae. In order to move sand and gravel, they shovel it with their snout and carry it in their mouths.
Preferred habitats vary with the season, temperature, dissolved oxygen content, flow
rate of the water, and the precipitation.
- Habitat Regions
- temperate
- freshwater
- Aquatic Biomes
- rivers and streams
Physical Description
Hatchlings have rounded heads, compared to the square, elongated heads of the adults. The mean length of hatchlings is estimated to be 22.8 mm long. They feature dark, well-formed eyes and developed forelimbs. Often on the forelimbs, three of the toes are complete and the fourth, inner toe is only a bud. The hindlimbs may not be fully developed and pressed close to the lower tail fin. They are light brown in color with dark lines from each nostril to the gills, and a white patch (approximately the size of their eye) behind and above the eye. They have melanophores on the gills, upper surfaces of the legs, lower jaw, and parts of the head, with concentrations highest in the area of the tail.
Studies have found that about 48% of the post-hatchling larvae have a dark lateral stripe while the rest have dark, poorly defined sides. Most have a broad, light tan, dorsal stripe from the snout to the tail. Along the dorsal region are small, dark (but poorly defined) spots. The underside of the larvae is white, or has a faint network of lines. After a year, the larvae measure 30 mm in length.
Adults lose the striped pattern; the side melanophores decrease in intensity while the dorsal melanophores increase in intensity and definition. The melanophores are dark, while the rest of the skin is reddish brown. Individuals with a SVL (snout vent length, measured from the tip of the snout to the vent) of 63 mm are considered adults. Adults have a set of bushy, red gills (gills are present in juvenile forms as well), as well as a set of weak lungs. Their tail is laterally compressed and they lack the fins present in the hatchlings. Adults have four toes on each foot. Neuse River waterdogs contain low levels of toxins in their skin to deter predators.
A major difference between the larvae of
N. lewisi
and other members of the genus
Necturus
is the difference in pigmentation.
Necturus maculosus
contains a dark dorsal area bordered by a thin, light, lateral stripe, while
Necturus punctatus
are uniformly gray with no striping pattern. The difference in the larvae adds to
the evidence that
N. lewisi
is a distinct species.
- Other Physical Features
- ectothermic
- heterothermic
- bilateral symmetry
- poisonous
- Sexual Dimorphism
- sexes alike
Development
Neuse River waterdogs hatch from eggs between April and July. Hatchling development
lags on their hindlimbs and on a fourth toe on their forelimbs. During the transition
from hatchling to larvae, the animals lose their tail fins and tadpole-like belly
and develop fully formed hindlimbs. The transition from larvae to adult includes change
in distribution of melanophores, and therefore, coloration, as well as an increase
in size and length. After 5.5 to 6.5 years, the waterdog is sexually mature and capable
of reproduction. Waterdogs can reach a total length of 23 cm and retain the use of
their gills as adults.
- Development - Life Cycle
- neotenic/paedomorphic
- metamorphosis
Reproduction
Courtship between males and females begins near the female's home retreat and lasts for about 1 hour. The male follows the female around, roughly 2 to 4 cm from her tail. When the female stops moving, the male crawls so its snout is behind a hindlimb of the female. At this time, the female's gills flare, while the male's retract. While the female is motionless, the male moves across the female's body and strokes her with his chin. Now, both the male's and female's gills flare and pulsate. The male then circles the female, while maintaining contact with her body, finally lying parallel to the female.
Males and females mate during fall and winter. During mating season, the male's vent
inflames. Males contain two papillae that enable the deposition of spermatophores
into the female's spermathecae, which hold the sperm until the following spring. After
eggs are laid, both parents guard the nest.
- Mating System
- monogamous
Neuse waterdogs reach sexual maturity at around 5.5 to 6.5 years or at a SVL (snout
vent length, measured from the tip of the snout to the vent) of 102 mm for males and
100 mm for females. Breeding once per year, males and females mate during fall and
winter and spawn in the spring. After displays of courtship, males deposit a packet
of sperm, which the female then places into her vent. Egg fertilization occurs internally
and during the spring, females will lay a clutch of eggs and both the male and female
will guard the nest. Females lay between 25 and 90 eggs.
- Key Reproductive Features
- iteroparous
- seasonal breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- fertilization
- oviparous
- sperm-storing
- delayed fertilization
After displays of courtship during the mating period, females allow males into their
home retreat area. During the spring, when the pair creates a nest for the eggs, both
sexes guard the nest until the young hatch. Because of the large clutch size, gamete
production is the most significant parental investment.
- Parental Investment
- male parental care
- female parental care
-
pre-fertilization
- provisioning
-
protecting
- female
-
pre-hatching/birth
-
provisioning
- female
-
protecting
- male
- female
-
provisioning
Lifespan/Longevity
Research has not been conducted on the longevity of
Necturus lewisi.
However, its relative
Necturus maculosus,
which resides in the Central United States and Canada, may live for 30 or more years.
Behavior
Home retreat areas are developed by waterdogs within 24 hours of exposure to a new environment. They push and carry gravel around to create an oval shaped area, usually with a diameter two-thirds the size of the animal. The opening of the area is often on the downstream side of the rock or covering.
In a study in 1985, waterdogs were presented with algae covered rocks, which were cleaned within 48 hours and converted into a home retreat area.
Nuese River waterdogs are active at night, and leave their retreats in search of food.
There is no evidence of migrational movements in this species.
- Key Behaviors
- natatorial
- nocturnal
- motile
- solitary
- territorial
Home Range
Home ranges of waterdogs often overlap, regardless of the sex or season. Males have
a significantly larger home range than females; 73 square meters for males compared
with 17 square meters for females.
Communication and Perception
Waterdogs defend their territories from intruders by flaring their gills. If an intruder advances, the flared gills pulsate and the waterdog curls its upper lip. During a confrontation, false attacks are displayed before actual contact is made. Bites serve as warnings which usually cause little damage to the intruder. Seventy-five percent of bites occur at the base of the tail.
Olfactory cues also protect territory. Areas with the scent of another animal, even if the animal is not physically present, are strongly avoided by intruders.
Stalking prey involves both visual and olfactory senses.
The observation that males rub the females with their chins during courtship suggest
that pheromones may play a role in communication during breeding.
- Other Communication Modes
- pheromones
- scent marks
Food Habits
Animals from the genus Necturus eat crayfish , insect larvae and nymphs, crustaceans, mollusks, plant remains, and worms.
Seventy-four percent of the diet of adult Necturus lewisi consists of arthropods. The waterdogs also eat mollusks, crayfish, shrimp, earthworms, Plecoptera nymphs , Odonata naiads , and some small fish including darters and pirate perch . On one occasion, scientists discovered an adult worm snake ( Carphophis amoenus ) inside the salamander. However, scientists noted that only the tail showed signs of digestion.
The larvae, which are often not large enough to consume some aquatic organisms, eat arthropods (approximately 99% of their diet), including ostracods and copepods .
The lack of Ceratopogonidae in the larvae N. lewisi diet distinguish N. lewisi from N. punctatus , its close relative. The adult N. punctatus do not eat vertebrates while the adult N. lewisi do.
Since the diet varies from season to season, based on prey availability, Neuse River
waterdogs are an opportunistic feeder. Hunting for their prey requires both sight
and olfactory senses, which they use to discern movement from a meter away. When hunting,
which usually occurs at night, waterdogs stalk the prey, then stop within striking
distance (approximately 2 to 3 cm). If the prey moves, the waterdog attacks and swallows
the prey. Larger prey are sometimes regurgitated and then re-swallowed. All prey are
ingested whole.
- Primary Diet
-
carnivore
- eats non-insect arthropods
- Animal Foods
- amphibians
- fish
- insects
- terrestrial non-insect arthropods
- mollusks
- terrestrial worms
Predation
While waterdogs eat small fish, larger fish prey on waterdogs. In order to avoid predators,
waterdogs avoid hunting and leaving the cover of their home retreat area until nightfall.
They also have toxins in their skin to deter predators. Their cryptic coloration likely
aids in avoiding predators as well.
- Anti-predator Adaptations
- cryptic
Ecosystem Roles
Neuse River waterdogs are predators on many organisms including mollusks, crayfish, shrimp, earthworms, insects, and some small fish. The prey are found in a variety of areas, including flowing water, slack water, leaf beds, under logs and rocks, and in sand and gravel.
To avoid predators, such as fish, the waterdogs hunt for food at night.
Studies have shown that when
Necturus punctatus
occupies the same tank as
N. lewisi
,
N. lewisi
will attack and eat its smaller relative.
Economic Importance for Humans: Positive
Small and easy to care for, the genus
Necturus
is often used in research and education. Since it represents a rather simple tetrapod,
it is often used for teaching anatomy. It is also a textbook example of paedomorphosis.
Waterdog ancestors probably had well-developed lungs; now they require gills to respire.
- Positive Impacts
- research and education
Economic Importance for Humans: Negative
There are no documented adverse effects of Necturus lewisi on humans.
Neuse River waterdogs do contain toxins within their skin, however, levels of the
toxin are not enough to affect humans. Some believe Neuse River waterdogs bring bad
luck.
Conservation Status
Because they require clean, flowing water and larger streams, Neuse River waterdogs
are listed as a species of "special concern" in North Carolina. Destruction of its
habitat through the creation of dams and pollution of the rivers threaten this species.
Other Comments
Necturus lewisi
was first recognized as a distinct subspecies in 1924 by C.S. Brimley. He named the
animal after Frank B. Lewis, who provided him with many specimens.
Additional Links
Contributors
Cynthia Lai (author), University of Michigan-Ann Arbor, Phil Myers (editor), University of Michigan-Ann Arbor, Rachelle Sterling (editor), Special Projects.
- Nearctic
-
living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- temperate
-
that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).
- freshwater
-
mainly lives in water that is not salty.
- ectothermic
-
animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature
- heterothermic
-
having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.
- bilateral symmetry
-
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
- poisonous
-
an animal which has a substance capable of killing, injuring, or impairing other animals through its chemical action (for example, the skin of poison dart frogs).
- metamorphosis
-
A large change in the shape or structure of an animal that happens as the animal grows. In insects, "incomplete metamorphosis" is when young animals are similar to adults and change gradually into the adult form, and "complete metamorphosis" is when there is a profound change between larval and adult forms. Butterflies have complete metamorphosis, grasshoppers have incomplete metamorphosis.
- monogamous
-
Having one mate at a time.
- iteroparous
-
offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).
- seasonal breeding
-
breeding is confined to a particular season
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- fertilization
-
union of egg and spermatozoan
- internal fertilization
-
fertilization takes place within the female's body
- oviparous
-
reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.
- sperm-storing
-
mature spermatozoa are stored by females following copulation. Male sperm storage also occurs, as sperm are retained in the male epididymes (in mammals) for a period that can, in some cases, extend over several weeks or more, but here we use the term to refer only to sperm storage by females.
- delayed fertilization
-
a substantial delay (longer than the minimum time required for sperm to travel to the egg) takes place between copulation and fertilization, used to describe female sperm storage.
- male parental care
-
parental care is carried out by males
- female parental care
-
parental care is carried out by females
- natatorial
-
specialized for swimming
- nocturnal
-
active during the night
- motile
-
having the capacity to move from one place to another.
- solitary
-
lives alone
- territorial
-
defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- chemical
-
uses smells or other chemicals to communicate
- pheromones
-
chemicals released into air or water that are detected by and responded to by other animals of the same species
- scent marks
-
communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- chemical
-
uses smells or other chemicals to communicate
- cryptic
-
having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.
- carnivore
-
an animal that mainly eats meat
References
Adams, L. 1926. Necturus: A Laboratory Manual . New York: J. J. Little and Ives Company.
Ashton, R., A. Braswell. 1979. Nest and Larbae of the Neuse River Waterdog, Necturus lewisi (Brimley) (Amphibia: Proetidae). Brimleyana , 1: 15-22.
Ashton, R. 1985. Field and Laboratory Observations on Microhabitat Selection, Movements, and Home Range of Necturus lewisi (Brimley). Brimleyana , 10: 83-106.
Brandon, R., J. Huheey. 1985. Salamander Skin Toxins, With Special Reference to Necturus lewisi. Brimleyana , 10: 75-82.
Braswell, A., R. Ashton. 1985. Distribution, Ecology, and Feeding Habits of Necturus-Lewisi (Brimley). Brimleyana , 10: 13-36.
Cooper, J., R. Ashton. 1985. The Necturus lewisi Study: Introduction, Selected Literature Review, and Comments on the Hydrologic Units and Their Faunas. Brimleyana , 10: 1-12.
Cooper, J. 1979. The Brothers Brimley; North Carolina Naturalists. Brimleyana , 1: 1-14.
Gilbert, S. 1977. Pictorial Anatomy of the Necturus . Seattle: University of Washington Press.
McDaniel, T., P. Martin, G. Barrett, K. Hughes, A. Gendron, L. Shirose, C. Bishop. 2009. Relative Abundance, Age Structure, and Body Size in Mudpuppy Populations in Southwestern Ontario. Journal of Great Lakes Research , 35 (2): 182-189.
Petranka, J. 1998. Salamanders of the United States and Canada . Washington: Smithsonian Institute Press.
Pudney, J., J. Canick, G. Callard. 1985. The Testis and Reproduction in Male Necturus, with Emphasis on N. lewisi (Brimley). Brimleyana , 10: 53-74.
United States Department of the Interior Fish and Wildlife Service. Conservation of the Amphibia of the United States: A Review. 134. Washington D.C.: United States, Fish and Wildlife Services. 1980. Accessed April 03, 2010 at http://www.nwhc.usgs.gov/disease_information/amphibian_malformation_and_decline/amphibian_conservation_1980.pdf .