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Necturus

Necturus is a genus of fully aquatic salamanders belonging to the family Proteidae, characterized by their neotenic development, retaining external gills and larval features throughout adulthood.^[1] Endemic to the eastern United States and Canada, species in this genus inhabit rivers, lakes, and streams, where they lead a secretive, bottom-dwelling lifestyle.^[2] Commonly referred to as mudpuppies or waterdogs, Necturus species are distinguished by their bushy, red or maroon external gills, four-toed feet, and laterally compressed tails adapted for swimming.^[3] The genus currently comprises seven recognized species: N. alabamensis (Alabama waterdog), N. beyeri (Gulf Coast waterdog), N. lewisi (Neuse River waterdog), N. maculosus (common mudpuppy), N. moleri (Moler waterdog), N. mounti (Ouachita waterdog), and N. punctatus (dwarf waterdog).^[1] These salamanders vary in size, with the largest, N. maculosus, reaching up to 49 cm in total length, while smaller species like N. punctatus typically measure 10–20 cm.^[4] Their skin is typically brownish-gray to black, often with dark spots or blotches, providing camouflage in muddy substrates.^[5] All Necturus species are obligate aquatics, possessing small rudimentary lungs but relying primarily on gill respiration, and they can also absorb oxygen through their skin. Necturus salamanders are carnivorous, feeding primarily on aquatic invertebrates such as crayfish, insects, and snails, as well as small fish and occasional plant matter.^[3] They are nocturnal and exhibit slow locomotion, using their broad heads and labial folds to suction-feed on prey.^[2] Reproduction occurs externally, with courtship in autumn or winter and egg-laying in spring; females deposit clusters of 50–200 eggs under rocks or logs, often guarded by one parent.^[2] Larvae hatch with gills and develop slowly, reaching maturity in 4–8 years depending on the species and environmental conditions.^[6] While most Necturus species are considered stable, some face threats from habitat degradation, water pollution, and collection for bait, leading to conservation concerns for taxa like N. alabamensis and N. lewisi.^[7] Their ecological role includes serving as indicators of water quality and as prey for larger aquatic predators.^[4] Ongoing research focuses on their phylogeny, with molecular studies refining species boundaries within the genus.^[1]

Taxonomy and Evolution

Taxonomy

Necturus is a genus within the family Proteidae, order Urodela (synonym Caudata), class Amphibia, comprising fully aquatic, paedomorphic salamanders characterized by persistent external gills in adults.^[1]^[8] The genus is divided into two subgenera: Necturus (for larger species) and Parvurus (for smaller coastal species such as N. punctatus).^[9] The etymology of the genus name derives from the Greek "nektos," meaning swimming, and "oura," meaning tail, reflecting the animal's fully aquatic lifestyle and possession of a laterally compressed tail for propulsion.^[4] The genus Necturus was first described by Constantine Samuel Rafinesque in 1818, based on specimens from the Ohio River drainage, with Necturus maculosus designated as the type species.^[10] Subsequent taxonomic revisions, such as Viosca's 1937 tentative classification, expanded recognition of species and subspecies within the genus, but ongoing debates persist regarding species boundaries, particularly in the N. beyeri complex where morphological variation and genetic divergence challenge the distinction of multiple lineages.^[11] Molecular phylogenetic studies confirm Necturus as the sister genus to Proteus within Proteidae, with the family occupying a basal position among the Cryptobranchoidea based on analyses of mitochondrial and nuclear DNA sequences.^[12]

Evolutionary History

The fossil record of Necturus spans from the late Paleocene to the present day, with all known fossils originating from North America and encompassing a temporal range of approximately 66 million years. The earliest undoubted proteid salamander, Necturus krausei, is documented from the late Paleocene Paskapoo Formation in Saskatchewan, Canada, based on well-preserved vertebrae that exhibit characteristic proteid features such as amphicoelous centra and neural arch morphology.^[13] Pre-Pleistocene records are sparse, limited primarily to these Paleocene vertebrae from the Western Interior Cretaceous Paleocene, while later Pleistocene fossils from sites like Florida indicate continuity into the Quaternary.^[14] Phylogenetic analyses of Necturus based on molecular genetic data, including mitochondrial and nuclear markers, support the recognition of 11 distinct lineages within the genus. A basal divergence separates the clade containing the Atlantic Coastal Plain species N. lewisi and N. punctatus from a more diverse assemblage of nine lineages restricted to Gulf Coastal Plain drainages. This east-to-west phylogeographic progression among lineages aligns with historical drainage patterns and underscores the role of riverine isolation in driving diversification. Paedomorphosis in the Proteidae, the family encompassing Necturus, is characterized by the obligate retention of larval traits—including external gills, lateral line organs, and a fully aquatic lifestyle—into sexual maturity, marking a derived evolutionary adaptation relative to the metamorphic life cycle of ancestral lissamphibians. Unlike the facultative paedomorphosis seen in some other salamander groups, this condition in proteids arose through heterochronic shifts that decoupled somatic maturation from reproductive development, facilitating specialization to permanent aquatic habitats.^[15] Such paedomorphic evolution has occurred independently in multiple urodele lineages but is fixed in Proteidae, contrasting with the terrestrial post-metamorphic phase typical of early amphibians. Genome size in salamanders underwent a significant increase in the ancestral Urodela, contributing to the neotenic (paedomorphic) traits observed in Necturus. With haploid genome sizes ranging from 80 to 95 pg in N. maculosus—among the largest in vertebrates—this expansion, driven by retrotransposon proliferation and intron lengthening, slows cell differentiation and developmental rates, thereby promoting the retention of juvenile morphology and physiology.^[16]^[17]

Diversity and Distribution

Species

The genus Necturus currently includes seven recognized species of fully aquatic, neotenic salamanders endemic to eastern North America.^[1] Taxonomic revisions based on molecular data suggest additional cryptic diversity within some lineages.^[18] These species are primarily distinguished by variations in adult size, body proportions, coloration patterns, and gill morphology, with N. maculosus representing the largest form, attaining lengths up to 49 cm.^[6] Historical synonyms reflect early taxonomic confusion, often stemming from regional descriptions and limited morphological data. Necturus alabamensis Viosca, 1937, known as the Alabama waterdog, is a medium-sized species (up to 24 cm) with a robust body, dark dorsal blotches on a grayish background, and relatively short limbs; it was originally described from the Black Warrior River system and has no major synonyms.^[19] Necturus beyeri Viosca, 1937, the Gulf Coast waterdog, reaches about 20 cm and features a slender build, uniform grayish-brown dorsum with faint spots, and prominent bushy gills; synonyms include N. lodingi Viosca, 1937, which was initially proposed as a distinct form but later synonymized, though recent analyses indicate potential separation.^[20]^[18] Necturus lewisi Trauth, 1993, or Neuse River waterdog, is similar in size to N. beyeri (up to 21 cm) but distinguished by a more mottled pattern and narrower head; it lacks notable historical synonyms, having been described relatively recently from North Carolina drainages.^[21] Necturus maculosus (Rafinesque, 1818), the common mudpuppy, is the most widespread and largest species (up to 49 cm), characterized by a broad head, rusty-brown body with irregular black spots, and highly branched red gills; original synonyms include Sirena maculosa Rafinesque, 1818, and Menobranchus lateralis Say, 1825. Some populations, such as those in the Red River, were formerly classified as a subspecies (N. m. louisianensis Viosca, 1938) but recent studies propose full species status (N. louisianensis) based on genetic and morphological differences.^[6]^[10]^[22] Necturus moleri Means and Trauth, 2020, the Moler waterdog, is a small to medium-sized species (up to 20 cm) with a slender body, mottled brown coloration, and moderately branched gills; it was recently described from the Apalachicola River basin.^[23] Necturus mounti Means, Lamb, and Long, 1979, the Ouachita waterdog, reaches up to 25 cm with a robust build, dark gray to black dorsum with light spots, and bushy gills; it is named after its type locality in the Ouachita River system.^[24] Necturus punctatus (Gibbes, 1850), the dwarf waterdog, is the smallest (up to 18 cm) with a delicate build, pale grayish coloration, fine spotting, and reduced gill branching; historical synonyms encompass Menobranchus punctatus Gibbes, 1850, and N. sorberi Viosca, 1937.^[25] Taxonomic uncertainties persist, particularly regarding subspecies like N. m. louisianensis and potential splits in N. beyeri (e.g., N. lodingi) and N. maculosus, where mitochondrial DNA studies have identified up to 11 distinct lineages warranting further investigation for full species recognition.^[26]

Geographic Distribution

The genus Necturus is endemic to eastern North America, with species distributed from southeastern Canada southward to the Gulf Coastal Plain, primarily in riverine and lacustrine systems east of the Great Plains.^[1] This range encompasses diverse physiographic regions, including the Great Lakes basin, Appalachian highlands, and Atlantic and Gulf coastal lowlands, but excludes western expansions beyond the Mississippi River drainage in most cases.^[27] Among the species, N. maculosus exhibits the broadest distribution, extending from the Great Lakes region and southeastern Canada (including parts of Ontario and Quebec) southward to Louisiana and Georgia, and westward to eastern Oklahoma and North Dakota.^[6] Several Necturus species display high endemism, confined to specific river basins that reflect localized evolutionary divergence. For instance, N. alabamensis is restricted to the Black Warrior River system in northwestern Alabama, where it occupies tributaries of the Tombigbee River.^[19] Similarly, N. lewisi is endemic to the Neuse and Tar-Pamlico River basins in the Piedmont and Coastal Plain of North Carolina, with no confirmed occurrences outside this narrow area.^[21] Other endemics include N. mounti, limited to the Choctawhatchee and Chipola River drainages in Alabama and the western Florida Panhandle, and N. moleri, found exclusively in the Apalachicola-Chattahoochee-Flint River basin spanning Georgia, Alabama, and Florida.^[24] These restricted ranges contrast with more widespread species like N. punctatus, which occurs along the Atlantic Coastal Plain from southeastern Virginia to central Georgia.^[25] Historically, Necturus distributions were more continuous across unglaciated refugia in the southeastern United States, but current ranges have contracted due to habitat fragmentation from anthropogenic alterations such as dams and channelization, particularly in northern populations of N. maculosus.^[28] No evidence supports transatlantic dispersal or westward expansions beyond historical limits, as the genus remains confined to North American freshwater systems.^[27] Biogeographic patterns within the genus are strongly influenced by Pleistocene glaciation, which isolated populations in southern refugia and drove post-glacial diversification and recolonization northward, leading to genetic structuring observed in species like N. maculosus.^[29]

Description

Physical Characteristics

Necturus species exhibit paedomorphic morphology, retaining larval characteristics throughout adulthood, such as persistent external gills and a fully aquatic lifestyle. Adults in the genus typically measure 20–25 cm in total length, though the common mudpuppy (N. maculosus) can reach up to 49 cm, making it the largest member of the group.^[6]^[21] Externally, these salamanders feature bushy, red or maroon external gills arising from two gill slits behind the head, which serve as the primary respiratory organs. The skin is smooth and glandular, lacking scales or eyelids, while the head is broad and flattened with small eyes. Limbs are short and stout, bearing four toes on both front and hind feet, and the tail is laterally compressed with dorsal and ventral fins, facilitating propulsion in water.^[6]^[30]^[31] Internally, Necturus lacks functional lungs, relying almost entirely on gill and cutaneous respiration for oxygen exchange, though rudimentary lung structures are present but seldom used. The vertebral column includes approximately 18 precaudal vertebrae, supporting a robust trunk suited to an aquatic existence.^[32]^[33] Sexual dimorphism is subtle but evident during the breeding season, when males develop a swollen cloaca with enlarged papillae to facilitate sperm transfer, while females retain a simpler cloacal structure.^[21]^[34]

Adaptations

Necturus species exhibit specialized respiratory adaptations that support their obligate aquatic lifestyle in often hypoxic environments. Their hemoglobin displays exceptionally high oxygen affinity, with a P50 value of 10.6 mmHg at 20°C—the highest recorded among salamanders—enabling efficient oxygen uptake from oxygen-poor water.^[35] This property is modulated by factors such as pH, temperature, and ATP, where ATP not only allosterically reduces oxygen affinity but also diminishes its temperature sensitivity, minimizing arterio-venous oxygen unloading differences across thermal gradients.^[35] Complementing this, the bushy external gills provide a large surface area for gas exchange, with individuals in low-oxygen habitats developing longer and more extensive gill structures to enhance extraction efficiency.^[36] These gills, retained through neoteny, facilitate predominant branchial respiration, supplemented minimally by cutaneous and pulmonary pathways.^[32] Sensory adaptations in Necturus prioritize aquatic perception over terrestrial vision. The eyes are small and reduced, lacking eyelids and positioned beneath a thin layer of skin, which limits visual acuity but suits their permanent submersion in murky waters.^[37] In contrast, the lateral line system is highly developed, featuring mechanoreceptive neuromasts along the head, body, and tail that detect water vibrations, pressure changes, and low-frequency movements with high sensitivity.^[37] This system, innervated by cranial nerves, generates receptor potentials in hair cells with amplitudes less than 800 microvolts in response to mechanical stimuli, aiding in navigation, prey detection, and predator avoidance in low-visibility habitats.^[38] Locomotor adaptations enable effective movement across benthic and pelagic zones. The feet bear four toes each, with moderate webbing that provides propulsion during swimming while allowing stable walking on substrates.^[6] The tail is laterally compressed with a dorsal and ventral fin, functioning as a primary sculling organ for efficient undulatory swimming.^[39] The bushy external gills, beyond respiration, contribute to hydrodynamic stability by increasing body surface area and aiding in gentle propulsion through water currents.^[40] Neoteny in Necturus manifests as the permanent retention of larval traits, including external gills and a tail fin. These features persist due to heterochronic shifts that prevent metamorphosis.^[41]

Reproduction and Life Cycle

Reproduction

Reproduction in Necturus species occurs annually, with courtship typically beginning in the fall or late winter and extending into early spring, often triggered by rising water temperatures that signal the transition to breeding conditions.^[42]^[43] In N. maculosus, for example, males initiate interactions by swimming or crawling around receptive females in shallow aquatic habitats, positioning themselves to deposit a spermatophore—a gelatinous, stalk-like packet of sperm approximately 10–12 mm long and 6–8 mm wide—on the substrate.^[44]^[42] The female then aligns her cloaca over the spermatophore, drawing it into her reproductive tract for internal fertilization, after which the male provides no further involvement in reproduction.^[43]^[45] Egg deposition follows in late spring, from April to June depending on latitude and temperature, with females selecting sheltered sites such as the undersides of rocks, logs, or other cover objects in streams or rivers.^[42]^[44] Each female lays a clutch of 18–174 eggs, typically averaging 50–100, which are attached individually in a loose monolayer rather than dense clusters, forming nests up to 15–30 cm in diameter; the eggs are creamy yellow, 5–6.5 mm in diameter, and surrounded by three jelly envelopes that increase their total size to about 15 mm.^[42]^[46] Fertilized eggs develop over 6–9 weeks, hatching into aquatic larvae that remain in the nest vicinity.^[43] Females exhibit nest-guarding behavior, remaining in or near the nest to defend the eggs against predators and disturbances, often coiling around them or positioning their bodies to shield the clutch until hatching.^[45]^[42] This parental care is energetically costly, as females may forgo foraging, leading to observations of filial cannibalism where starved females consume some of their own eggs to sustain themselves.^[45] Such behavior has been documented in N. maculosus, marking it as one of few aquatic salamander genera showing this trait during nest attendance.^[45] While detailed for N. maculosus, reproductive patterns are similar across Necturus species, with minor variations in clutch size and timing.^[1]

Development

The development of Necturus species, such as the common mudpuppy (N. maculosus), exemplifies paedomorphosis, where larval traits are retained into adulthood without undergoing metamorphosis. Eggs are attached individually in a loose monolayer to submerged substrates like rocks or logs in spring, and incubation lasts 1 to 2 months, influenced by water temperature, typically ranging from 30 to 70 days.^[47]^[48]^[5] Upon hatching, larvae measure approximately 20 to 25 mm (about 1 cm) in length and possess prominent yellow yolk sacs for initial nourishment, along with fully developed external gills that enable immediate aquatic respiration.^[48]^[5] These yolk-sac larvae remain concealed under the nest cover, guarded by the female, for 6–8 weeks post-hatching until the yolk sac is absorbed, typically leaving the nest by late summer (e.g., end of August).^[6]^[45] Once the yolk sac is absorbed, the larvae emerge to feed on small invertebrates, including insect larvae and microcrustaceans, supporting their transition to independent foraging.^[47]^[45] Necturus exhibits direct paedomorphosis, bypassing metamorphosis entirely; individuals reach sexual maturity while retaining larval features such as external gills, a laterally compressed tail fin, and an entirely aquatic lifestyle indefinitely.^[49]^[50] This neotenic retention of juvenile morphology allows for reproduction without the typical amphibian transformation to a terrestrial form.^[51] Growth in Necturus is slow, with individuals reaching sexual maturity in 4 to 6 years at a length of about 20 cm, though some sources report maturity around 5 years at 15 to 20 cm.^[47]^[52] Lifespan varies by environment, averaging 11 years in the wild but extending up to 20 years or more in captivity under optimal conditions.^[36]

Ecology and Behavior

Habitat and Diet

Necturus species are fully aquatic salamanders that inhabit permanent freshwater environments, including clear streams, rivers, and lakes with rocky or gravel substrates. They prefer well-oxygenated waters with moderate flow and temperatures typically ranging from 9 to 20°C, avoiding heavily stagnant or silty conditions that reduce visibility and shelter availability.^[53]^[6] These habitats provide ample cover, such as submerged rocks, logs, and vegetation, where individuals seek refuge during the day to avoid predation and desiccation.^[47] Microhabitat use is closely tied to their nocturnal lifestyle; Necturus are primarily active at night, foraging along the bottom in crevices or open substrates at depths of 0.5 to 2 meters, though they may occupy deeper waters up to 27 meters in lakes during certain seasons. Diurnally, they retreat to tight shelters like rock fissures or undercut banks, emerging primarily under low-light conditions. Seasonal patterns influence habitat selection, with individuals shifting to deeper, cooler waters in summer to avoid high temperatures and potentially reducing activity during extreme cold, though they remain active year-round in temperate regions.^[53]^[6]^[47] As opportunistic carnivores, Necturus feed on a variety of benthic invertebrates and vertebrates, with crayfish forming a significant portion of the diet (up to 33% by frequency in some populations), alongside insects (up to 92% frequency), annelids (up to 34% frequency), small fish (up to 22% volume or 13% frequency), and mollusks (up to 22% frequency).^[54]^[45] They employ suction feeding to capture prey, using rapid mouth expansion to draw in small items whole or larger ones through repeated gulps, detected via lateral line organs and olfaction rather than keen vision. Diet composition varies seasonally, with invertebrates dominating in summer and fall (e.g., higher insect intake) and fish more prevalent in winter and spring; foraging intensity may decrease in warmer months, leading to more empty stomachs.^[54]^[45]^[55]

Predators and Interactions

Necturus species face predation from a variety of aquatic and riparian predators, including larger fish such as bass (Micropterus spp.) and catfish (Ictalurus spp.), wading birds like great blue herons (Ardea herodias), and mammals including river otters (Lontra canadensis).^[56]^[43]^[5] Their prominent external gills, retained throughout life, increase vulnerability by providing an easily accessible target for attacks and potentially hindering escape from grasping predators.^[47]^[57] To counter these threats, Necturus exhibit primarily behavioral defenses, remaining nocturnal to minimize detection by diurnal predators and foraging under cover of darkness.^[53]^[58] Tail autotomy serves as a morphological escape mechanism, allowing individuals to shed and regenerate their tail if grasped, though this is less frequently observed compared to other salamanders.^[59] Additionally, skin secretions provide chemical deterrence, releasing noxious compounds that can repel potential attackers, a common antipredator strategy among amphibians.^[60] Symbiotic relationships in Necturus include serving as the exclusive host for the glochidia (larval stage) of the salamander mussel (Simpsonaias ambigua), which attach to the external gills without causing significant harm to the salamander.^[61]^[43] However, they also host various parasites, such as acanthocephalans, trematodes, and nematodes, which can infest the gills and intestines, potentially impacting health under high loads.^[62]^[63] Intraspecific interactions are generally limited in aggression, with individuals exhibiting territorial behaviors primarily during nesting, where males guard egg sites through displays such as foreleg circumduction to deter rivals.^[64] Cannibalism among larger individuals occurs occasionally but is not a dominant interaction.^[43]

Conservation

Status

The genus Necturus comprises seven species of fully aquatic salamanders endemic to eastern North America, with most facing conservation concerns due to their restricted geographic ranges and vulnerability to habitat degradation.^[65] While the common mudpuppy (N. maculosus) is assessed as Least Concern globally owing to its broad distribution across multiple watersheds, it is locally vulnerable in fragmented populations where water quality declines. Individual species statuses vary under the IUCN Red List, reflecting differences in range extent and threat exposure as of assessments through 2025. The Alabama waterdog (N. alabamensis) is Endangered due to severe habitat loss and pollution in its limited Black Warrior River basin range. The Neuse River waterdog (N. lewisi) is Near Threatened, with ongoing risks from river impoundments and sedimentation in North Carolina drainages. The dwarf waterdog (N. punctatus) is Least Concern, benefiting from a wider coastal plain distribution, though local declines are noted.^[66] The Gulf Coast waterdog (N. beyeri) is also Least Concern, with stable populations in Gulf drainages. Recently described species like the Escambia waterdog (N. mounti) are Not Evaluated, reflecting the need for further assessment in narrow river systems of Alabama and Florida. The Apalachicola waterdog (N. moleri) is Not Evaluated pending further surveys in its southeastern river habitats.^[23] Population trends differ markedly across the genus, with widespread species like N. maculosus showing overall stability but localized declines of up to 30-50% in polluted urban-adjacent waters over the past two decades. Endemic species exhibit sharper reductions; for instance, N. alabamensis has lost approximately 50% of its historical range since the 1980s due to coal mining runoff and channelization, leading to estimated population decreases exceeding 50% in occupied streams. Similarly, N. lewisi populations have declined by 20-40% in monitored Tar and Neuse River segments since 2000, driven by water quality deterioration. Several Necturus species receive legal protections under the U.S. Endangered Species Act (ESA), enhancing regulatory safeguards against habitat alteration. N. alabamensis has been listed as Endangered since 2018, prohibiting take and mandating habitat consultations in the Black Warrior basin.^[67] N. lewisi was designated Threatened in 2021, with a 4(d) rule allowing limited activities while protecting core river habitats in North Carolina.^[57] These listings apply wherever the species occur, supporting recovery efforts like water quality monitoring and restoration, including a September 2025 recovery plan for N. alabamensis outlining delisting criteria.^[68]^[69]

Threats and Protection

Necturus species face significant threats from habitat loss primarily caused by the construction of dams, stream channelization, and increased siltation from erosion and land-use changes, which degrade the rocky, well-oxygenated substrates essential for their survival.^[70]^[39] Pollution exacerbates these issues, with agricultural, industrial, and forestry runoff introducing heavy metals, organochlorine compounds, and other contaminants that accumulate in tissues and disrupt physiological functions.^[71]^[39] Lampricides used to control invasive sea lampreys pose acute risks, as these chemicals are toxic to non-target amphibians like mudpuppies (Necturus maculosus), leading to direct mortality during treatments.^[39]^[72] Additionally, bycatch in recreational and commercial fisheries, particularly through setlines, traps, and gill nets, results in unintended capture and high mortality rates for these gill-breathing salamanders.^[73]^[74] Climate change compounds these pressures by warming aquatic habitats, which reduces dissolved oxygen levels and exceeds the thermal tolerances of Necturus species adapted to cool waters, potentially shifting their distributions or causing population declines.^[43]^[75] Invasive species, such as zebra and quagga mussels (Dreissena spp.), further alter food webs by filtering plankton and competing for resources, indirectly threatening Necturus through reduced prey availability and habitat modification in the Great Lakes and connected rivers.^[76] Conservation efforts include habitat restoration projects in priority river systems, such as stabilizing streambanks and removing low-head dams to improve connectivity and water quality for Necturus populations. Efforts to protect symbiotic relationships, particularly the endangered salamander mussel (Simpsonaias ambigua), which relies exclusively on mudpuppies as hosts for its larval stage, involve mussel reintroduction and host population monitoring to ensure mutual survival.^[77] Ongoing monitoring programs, such as those outlined in New York's 2025 State Wildlife Action Plan (SWAP), track distribution and abundance through targeted surveys to inform adaptive management.^[78] Research priorities emphasize updated field surveys for data-deficient Necturus taxa to fill knowledge gaps on population trends and distributions. Genetic studies are also critical to assess diversity across fragmented populations, supporting the development of captive breeding protocols, as demonstrated in successful hormone-assisted reproduction for species like the Gulf Coast waterdog (N. beyeri).^[79]

References

[1]
Proteidae - AmphibiaWeb
One species of Proteus and five species of Necturus are currently recognized. The Olm, or European blind cave salamander, Proteus anguinus, is perhaps the ...
[2]
Proteidae (Olms, Waterdogs, Mudpuppies) - Animal Diversity Web
Four Necturus species, the waterdogs, grow to 28 cm in length, and the fifth, N. maculosus , the mudpuppy, grows to more than 40 cm long. Their common names ...<|separator|>
[3]
Necturus maculosus (Common Mudpuppy) - DigiMorph
Sep 23, 2003 · Necturus is a perennibranchiate (retaining external gills) salamander of the Euramerican clade Proteidae. The genus is confined to North ...Missing: biology | Show results with:biology
[4]
Mudpuppy - INHS Herpetology Collection
Description: Large (up to 34 cm TL), stout-bodied brownish gray, rust brown, or black salamander with scattered round black spots of various sizes.
[5]
Common Mudpuppy - Missouri Department of Conservation
All mudpuppy species (genus Necturus) have adults that are neotenic: they retain larval characteristics, namely the gills, into adulthood. Similar species ...
[6]
Necturus maculosus - AmphibiaWeb
Description Completely aquatic and gilled throughout life. The mudpuppy is the largest member of the genus Necturus which also includes waterdogs.
[7]
Necturus - Explore the Taxonomic Tree | FWS.gov
Necturus punctatus · Dwarf Waterdog · 173625 ; Necturus lewisi · Neuse River Waterdog · 173627 ; Necturus alabamensis · Black Warrior Waterdog, Alabama Waterdog, Black ...
[8]
Proteidae Bonaparte, 1831 | Amphibian Species of the World
Evol., 53: 492–508, reported on molecular phylogenetics of salamanders based on mtDNA and provided an estimate of time since origin of the salamander families.Missing: phylogeny | Show results with:phylogeny
[9]
Necturus maculosus (Rafinesque, 1818)
Necturus maculosus (Rafinesque, 1818). Class: Amphibia > Order: Caudata > Family: Proteidae > Genus: Necturus > Species: Necturus maculosus. Proteé ...
[10]
Evolutionary insights into the North American Necturus beyeri ...
Dec 13, 2017 · Our work suggests that Necturus comprises 11 lineages. A basal split within the genus separates an ancestor of two Atlantic Coastal Plain ...
[11]
Analyses of Mitochondrial and Nuclear Genomic Data | Systematic ...
To reexamine these relationships and the phylogenetic-information content of molecular characters, we present new mitochondrial DNA (mtDNA) sequences, ...
[12]
The Earliest Known Necturus (Amphibia, Urodela), from the ... - jstor
Necturus krausei represents the earliest known undoubted proteid, and is probably the only pre-Pleistocene record for the family in North America. A vomer ...
[13]
Biogeography and body size shuffling of aquatic salamander ...
The pre-Pleistocene fossil record of Necturus is limited to two small but well-ossified Palaeocene vertebrae from the WICP [39].
[14]
Paedomorphosis and Higher-Level Salamander Relationships
A recent study of higher-level salamander phylogeny placed most paedomorphic families in a single clade based on morphological data. Here, we use new molecular ...
[15]
Forever young: Linking regeneration and genome size in salamanders
Dec 15, 2020 · Here we show how to unite seemingly disparate facets of salamander biology, regeneration ability, and genome size variation, into one synthetic view.
[16]
[PDF] Repeated ecological and life cycle transitions make salamanders an ...
Apr 16, 2021 · Necturus maculosus has a haploid genome size of 80 to. 95 pg ... cycle evolution is linked with genome size in salamanders. (Figure 2) ...
[17]
Necturus Rafinesque, 1819 | Amphibian Species of the World
### Recognized Species in Genus Necturus
[18]
Colour and size reveal hidden diversity of Necturus (Caudata
The genus Necturus traditionally contains five species of paedomorphic salamanders restricted to the eastern United States (Highton et al. 2017).
[19]
Necturus alabamensis - AmphibiaWeb
family: Proteidae subfamily: Necturinae genus: Necturus · Necturus ... Among mudpuppies and waterdogs (genus Necturus), this species is medium-sized.
[20]
Necturus beyeri - AmphibiaWeb
This species is identified as a member of the family Proteidae by retention of external gills in adults, and the presence of four robust limbs with four ...
[21]
Necturus lewisi - AmphibiaWeb
Description Completely aquatic and gilled throughout life. Among mudpuppies and waterdogs (genus Necturus), this species is medium-sized.
[22]
Necturus louisianensis Viosca, 1938 - Amphibian Species of the World
Necturus louisianensis Viosca, 1938. Class: Amphibia > Order: Caudata > Family: Proteidae > Genus: Necturus > Species: Necturus ...Missing: AmphibiaWeb | Show results with:AmphibiaWeb
[23]
Necturus louisianensis - NatureServe Explorer
Amphibia. Order: Caudata. Family: Proteidae. Genus: Necturus. Scientific Name Reference: Frost, D.R. 2020. Amphibian Species of the World: an Online Reference.
[24]
(PDF) Colour and size reveal hidden diversity of Necturus (Caudata
Mar 19, 2020 · Recent data from the mitochondrial genome reveal six lineages of Gulf Coast Waterdogs traditionally classified as Necturus beyeri.
[25]
Co-occurrence and Hybridization between Necturus maculosus and ...
Oct 20, 2017 · The genus Necturus (Proteidae) includes five species of large neotenic salamanders commonly referred to as mudpuppies and waterdogs. All are ...Missing: boundaries | Show results with:boundaries
[26]
Necturus mounti - AmphibiaWeb
Aug 17, 2020 · This species is identified as a member of the family Proteidae by retention of external gills in adults, and the presence of four robust ...
[27]
Necturus punctatus - AmphibiaWeb
Description Completely aquatic and gilled throughout life. · Distribution and Habitat · Life History, Abundance, Activity, and Special Behaviors Little is known ...
[28]
Mudpuppy (Necturus maculosus maculosus ) spatial distribution ...
Mudpuppy (Necturus maculosus maculosus) populations have been declining in the Great Lakes region of North America.
[29]
Phylogeographic analysis of Mudpuppies (Necturus maculosus)
The geology of the Pleistocene, and particularly the Last Glacial Maximum approximately 26.5 ka, is a critical driver of species' present-day distributions ...
[30]
Common Mudpuppy - an overview | ScienceDirect Topics
This chapter examines the external and internal anatomy of the mudpuppy. The common mudpuppy, Necturus maculosus, is a member of Caudata, which together with ...
[31]
Mudpuppy | Necturus maculosus - Wisconsin DNR
Mudpuppies are 12-16 inches, Wisconsin's largest aquatic salamander with red gills, rusty-brown or grayish color, and live in lakes and rivers. They are not ...Missing: lanthami | Show results with:lanthami
[32]
[PDF] Gas exchange and habitat selection in the aquatic salamanders
In the case of Necturus, the gills have been presumed to allow it to occupy hypoxic waters in spite of its poor lungs, which may explain its much wider range ( ...
[33]
[PDF] Variations In The Vertebrae And Pelvis Of Necturus - SMU Scholar
Parker (1896) examined the skeletons of 27 Necturus. Nineteen of the 27 specimens had Vertebra XIX as the sacrum, while six had Vertebra XX. In one specimen the.Missing: count | Show results with:count
[34]
Necturus maculosus (Mudpuppy)
The Mudpuppy is a large aquatic salamander with adult lengths of 20.3 to 48.2 cm (8-19 in). It has a broad, flat head with small eyes, a dark stripe through ...
[35]
Adaptations to neoteny in the salamander, Necturus maculosus ...
Adaptations to neoteny in the salamander, Necturus maculosus. Blood respiratory properties and interactive effects of pH, temperature and ATP on hemoglobin ...
[36]
[PDF] Mudpuppy Fact Sheet - CT.gov
The mudpuppy is recognized by red to maroon-colored bushy gills; a gray-green and mottled back with blue-black spots and a gray belly; a broad flattened head ...
[37]
Necturus - an overview | ScienceDirect Topics
Hynobiidae, the other cryptobranchoid lineage, has no fossil record. Proteids occurred first in the Late Paleocene of North America and the Middle Miocene of ...
[38]
Receptor Potentials From Hair Cells of the Lateral Line - PubMed
Intracellular recordings from hair cells in the tail lateral line of mudpuppy Necturus maculosus show receptor potentials less than 800 microvolts.Missing: adaptations eyes
[39]
Mudpuppy (Necturus maculosus) Manitoba population Great Lakes ...
Oct 21, 2024 · The Mudpuppy (Necturus maculosus) is a large aquatic salamander that ranges up to 49 cm in length. Its most obvious feature is prominent, ...
[40]
[PDF] The Natural History of Necturus: II - SMU Scholar
Necturus has permanent external gills, simple internal lungs, and a skin well supplied with blood vessels; all of which have been reported functional in ...Missing: adaptations sensory neoteny scientific
[41]
Repeated ecological and life cycle transitions make salamanders an ...
May 15, 2021 · This article summarizes ecological and endocrine-based causes of life cycle transitions in salamanders, as well as consequences to body size, genome size, and ...
[42]
Necturus maculosus - Amphibians of North Carolina
These are the Common Mudpuppy (N. m. maculosus) that is found throughout most of the range of the species, and the Red River Mudpuppy (N. m. louisianensis) that ...
[43]
[PDF] Mudpuppy Species Guidance Document - Natural Heritage
Necturus is the only known glochidial host of the Salamander. Mussel (Simpsonaias ambigua). The latter species is listed as endangered in Illinois and is ...<|control11|><|separator|>
[44]
Mudpuppy - North Carolina - Davidson College
Reproduction: Courtship consists of males waiting for females in shallow water and coaxing the female to move her cloaca over a sperm packet (spermatophore) ...<|control11|><|separator|>
[45]
[PDF] Common Mudpuppy (Necturus maculosus) distribution, diet and ...
Feb 2, 2021 · Their distribution covers an area from southeastern Canada to Georgia and Louisiana, representing the largest distribution of any fully aquatic ...
[46]
[PDF] The Natural History of Necturus, IV. Reproduction - SMU Scholar
Any brooding activity on the part of Necturus is probably incidental; her back may oc- casionally brush the eggs as they hang from the protecting object, but ...
[47]
Necturus maculosus (Mudpuppy) - Animal Diversity Web
Mudpuppies are between 20 and 33 cm in length. They are neotenic (permanent larvae) and retain large, maroon colored external gills throughout their life.
[48]
Necturus maculosus - Mudpuppy - Kansas Herpetofaunal Atlas
The Mudpuppy is a large, obligate aquatic salamander with a large flattened, square head, small eyes, and a pair of distinctive, red feathery gills on each ...
[49]
Common Mudpuppy (Necturus maculosus (Rafinesque, 1818)) in ...
Sep 26, 2022 · The Common Mudpuppy (Necturus maculosus (Rafinesque, 1818)) is found in many watersheds from eastern Canada to the southeastern United ...
[50]
Necturus - an overview | ScienceDirect Topics
Paedomorphosis in the plethodontid Typhlomolge can be explained by underdevelopment of the thyroid [104]; however, the endocrine complex appears functional in ...
[51]
Are paedomorphs actual larvae? - American Association for Anatomy
Feb 2, 2021 · The resulting sexually mature animals, called paedomorphs, retain anatomically larval characters (gills, tailfin, etc.) ... Necturus, and the cave ...Missing: retention | Show results with:retention
[52]
Pademorphosis: Meaning and Types | Amphibians
Some populations of Cryptobranchus alleganiensis, after complete metamorphosis, retain a single gill-slit that reflects the ancestral larval feature. ... Necturus ...
[53]
Mudpuppy (Necturus maculosus)
Mudpuppies become sexually mature at 4–6 years old and breed in the fall. Eggs are fertilized internally, like most other salamanders. The female releases 75– ...
[54]
Common mudpuppy - Wikipedia
The common mudpuppy (Necturus maculosus) is a species of salamander in the family Proteidae. It lives an entirely aquatic lifestyle in parts of North ...<|control11|><|separator|>
[55]
[PDF] The Natural History of Necturus: I. Habitats and Habits - SMU Scholar
He suggests that the robust- bodied N. m. maculosus and N. maculosus beyeri hide under flat rocks, etc.; and_ that the other species, being better adapted.<|control11|><|separator|>
[56]
[PDF] The Natural History of Necturus: III. Food and Feeding - SMU Scholar
Their approach to possible food is cautious.,:- If lateral line stimulation is rein- forced by proper olfactory or optic stimulus, a snapping response is ...Missing: adaptations | Show results with:adaptations
[57]
(PDF) Necturus maculosus (Mudpuppy): predation - ResearchGate
Jan 26, 2022 · Necturus maculosus (Mudpuppy): predation. December 2021; Herpetological ... tra canadensis (River Otter), Ardea herodias (Great Blue Heron),.
[58]
Neuse River Waterdog (Necturus lewisi) | U.S. Fish & Wildlife Service
As a narrow endemic, it is only found in the Neuse and Tar-Pamlico River Basins of North Carolina (NC). It lives in medium to large streams and rivers in the ...
[59]
[PDF] Abundance of Gulf Coast Waterdogs (Necturus beyeri) along Bayou ...
Jun 11, 2021 · Gulf Coast Waterdogs (Necturus beyeri) are permanently aquatic, nocturnal salamanders with both lungs and external gills and inhabit lotic ...
[60]
Necturus Maculosus - an overview | ScienceDirect Topics
Studies showed that morphologically similar paedomorphic salamanders (with external gills or gill slits, no eyelids, larval pigmentation, etc.) responded ...
[61]
Chemical Signaling in Amphibians - NCBI - NIH
Many salamanders release noxious skin secretions that function in predator defense. In red-legged salamanders (Plethodon shermani), antipredator secretions ...
[62]
Salamander Mussel (Simpsonaias ambigua) | U.S. Fish & Wildlife ...
The mudpuppy (Necturus maculosus), the only known host for salamander mussel ... The salamander mussel's larvae (called glochidia) develop on the gills ...
[63]
Necturus beyeri - AmphibiaWeb
Necturus beyeri is a neotenic species of aquatic salamander with adults reaching a standard length of up to 184.0 mm in males and 177.0mm in females. Average ...
[64]
Survey of the Helminth Parasites Infecting the Alabama Waterdog ...
This study was designed to provide the foundation for future work on the ecology and evolution of Necturus-parasite systems by providing vital parasite survey ...
[65]
(PDF) Nest Descriptions for the Mudpuppy (Necturus maculosus)
Necturus maculosus is a rarely seen and poorly understood permanently ... Literature suggests that parturition and nest-guarding in N. m ...
[66]
AmphibiaWeb Search
IUCN Red List Status, Vernacular Name, Family, Order. Necturus alabamensis · photos, Endangered (EN), Alabama Waterdog, Proteidae, Caudata. Necturus beyeri ...
[67]
https://ecos.fws.gov/ecp/species/5426
[68]
Necturus moleri - AmphibiaWeb
Aug 18, 2020 · This species is identified as a member of the family Proteidae by retention of external gills in adults, and the presence of four robust limbs ...
[69]
Black warrior (=Sipsey Fork) Waterdog (Necturus alabamensis)
The Black warrior Waterdog (Necturus alabamensis) is an endangered species, listed as such on 02-02-2018, and found wherever it is, including Alabama.
[70]
Threatened Species Status With Section 4(d) Rule for Neuse River ...
Jun 9, 2021 · We, the US Fish and Wildlife Service (Service), list two North Carolina species, the Carolina madtom ( Noturus furiosus) as endangered, and the Neuse River ...<|separator|>
[71]
Necturus maculosus : Mudpuppy | Rare Species Guide | Minnesota ...
This fully aquatic salamander is found in medium to large rivers and larger lakes within the Mississippi, Minnesota, Red, and St. Croix river basins.
[72]
[PDF] COSSARO Evaluation Report for Mudpuppy
Mudpuppy (Necturus maculosus) is classified as Special Concern in Ontario. Mudpuppy does not currently meet criteria to be considered Endangered or Threatened, ...
[73]
[PDF] Assessment of mudpuppy (Necturus maculosus) presence along the ...
Mar 15, 2019 · Mudpuppies (Necturus maculosus) are an elusive, fully aquatic salamander that occupy streams, lakes, and ponds in eastern North America.
[74]
[PDF] Observations of Mudpuppy (Necturus maculosus) bycatch in a ...
Apr 3, 2018 · Siltation and chemical pollutants. (Boninet al.1995; Matson 1998, 2005) are likely chron- ic threats to local populations, and bycatch in rec ...
[75]
[PDF] Mudpuppy (Necturus maculosus) Assessment Along the St. Clair ...
Monitoring of Mudpuppy health in correlation to surrounding land uses may be a useful tool for identifying major threats in the region. Furthermore, continued ...<|control11|><|separator|>
[76]
Vulnerability of amphibians to global warming - Nature
Mar 5, 2025 · Overall, our study contributes to the evidence that climate change is a mounting threat to amphibians and emphasizes the importance of limiting ...
[77]
Diets, population structure, and seasonal activity patterns of ...
Stomach contents obtained through gastric lavage included mollusks, leeches, insects, isopods, amphipods, crayfish, fish, a frog, and a juvenile conspecific.Missing: nocturnal | Show results with:nocturnal
[78]
[PDF] Sampling protocol for the freshwater mussel Simpsonaias ambigua ...
This species is unique in that it uses an aquatic salamander with external gills. (Necturus maculosus (Rafinesque, 1818), Mudpuppy), as a host instead of a fish ...Missing: symbiosis SWAP
[79]
| Northeast Fish and Wildlife Diversity
2025 SWAP Database. Home. Habitats. Taxa · States ... Mudpuppy. Necturus maculosus. Species Details in ... S101P332, C.8.1.0.0, Basic research and status monitoring.
[80]
(PDF) Successful Captive Breeding of Necturus beyeri through ...
Aug 6, 2025 · Successful captive breeding of Necturus beyeri through manipulation of environmental cues and exogenous hormone administration: a model for endangered necturus.Missing: deficient | Show results with:deficient