Lesser siren

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Lesser siren
Siren intermedia 2.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Amphibia
Order: Urodela
Family: Sirenidae
Genus: Siren
Species:
S. intermedia
Binomial name
Siren intermedia
Barnes, 1826

The lesser siren (Siren intermedia) is a species of aquatic salamander native to the eastern United States and northern Mexico. They are referred to by numerous common names, including two-legged eel, dwarf siren, and mud eel. The specific epithet intermedia denotes their intermediate size, between the greater siren, Siren lacertina, and the dwarf sirens, Pseudobranchus species.

Contents

Description

Lesser sirens have elongated bodies possessing only two limbs, a pair of four-toed legs located behind the base of the head, and range in length from 7 to 27 inches (17 – 69 cm). Unlike greater sirens, lesser sirens have less than 35 costal grooves. [2] Juvenile specimens have red to yellow banding on their heads and stripes running along the main length of the body, although these stripes are absent in Siren intermedia. Adults have plainer coloration; the dorsal side is typically olive green to blue-gray or black, while the ventral is usually light grey. Spotted patterns may also be present in adults. [3] In salamanders, sexual size dimorphism is usually female-biased, but in Siren intermedia, it’s the opposite. Males are the larger sex, and they also have significantly larger heads and enlarged masseter muscles than females. [4]

Lesser sirens are notably neotenic, and possess external gills throughout their lives. [3]

Taxonomy

Siren intermedia nettingi, the western lesser siren. Siren intermedia nettingi UMFS 2015 1.JPG
Siren intermedia nettingi, the western lesser siren.

Sources disagree on the number of subspecies within S. intermedia. The majority of the sources in the field agree there are at least two: an eastern and a western variety. Many sources also include a third subspecies, the Rio Grande lesser siren, S. i. texana, but researchers disagree whether the Rio Grande variety belongs as a lesser siren, within S. intermedia, or as a greater siren, within S. lacertina, and some others even consider it to be its own species, as S. texana. [5] [6]

Habitat and distribution

Habitat

Sirens are likely to be found in ponds near other intermittent wetlands. They usually inhabit swamps and ponds, and can survive in dry ponds for up to two years by a process called “aestivating”. Sirens are considered poor overland dispersers because they rarely leave the water, so their geographical distribution is largely determined by pond connectivity and biotic interactions. [7]

Lesser sirens are aquatic salamanders, but their ability to withstand factors like dehydration allows them to have good survival skills in their ephemeral habitats. Sirens are able to osmoregulate and produce a mucoid cocoon during aestivation, which greatly reduces dehydration and electrolyte stress. [8] Their small legs enable them to move on dry land for short periods of time. [9]

Geographic distribution

The lesser siren is found in the United States and Mexico, primarily from Virginia to Florida, west to Texas, ranging into northeastern Mexico as far as Veracruz, and north to Illinois and southwest Michigan. [1] [10] Multiple specimens have recently been rediscovered in Michigan after a 60-year absence. [11] [12] Lesser sirens only occur in permanent wetlands, whereas other salamander and frog species can occupy a range of dry and wetlands. Lesser sirens are top predators in permanent wetlands. [13] They can quickly colonize and become dominant consumers in new ponds such as those constructed by beavers. Their ability to aestivate and burrow in soft sediments and live in dense aquatic vegetation in shallow waters facilitates their dominance. In addition, the species’ high productivity, high fecundity, and rapid growth to early sexual maturity allows it reach a high density quickly and easily, and thus establish dominance in the habitat. [14]

Conservation

The lesser siren is quite common through most of its range, but rarely seen due to its secretive nature. Like almost all species of amphibian, their numbers are believed to be declining due to general reductions in water quality caused by agricultural pesticide and fertilizer runoff. They are frequently collected and used as bait for fishing.

The species was once believed to be extirpated from Michigan, [11] but has been recently rediscovered in limited numbers. [15] The S. i. texana subspecies is listed as a threatened species in Texas. They are listed as Least concern on the IUCN Red List and a species with "Special Protection" in Mexico. [1]

Physiology

Vision

Western Lesser Sirens (Siren intermedia nettingi) have poor eyesight. They rely on their other senses to forage, hunt, and survive. [13]

Chemical sense

Sirens rely on chemical cues to detect prey and not on visual cues. The use of chemical stimuli in the detection of prey and predators are common in amphibians. [16]

Respiration

All species in the sirenidae family are paedomorphic, eel-shaped salamanders that live in aquatic environments. They respire through their gills, lungs and skin, and survive well in hypoxic environments low in oxygen. Siren intermedia is unique among amphibians in its ability to construct a mucoid cocoon that slows down dehydration during aestivation, and the lesser siren can withstand long periods of food deprivation without ill effects. During aestivation, respiration slows down significantly, and gills atrophy over the next 16 or more weeks. [8]

Acoustic behavior

The lesser siren is vocal, unlike most salamanders. The lesser siren is known to emit a series of clicks when it approaches others of its species, or a short screeching sound if handled. Acoustic behavior serves a functional service in S. intermedia, especially since it is nocturnal and it burrows in sediments, swims and crawls through densely vegetated waters. However, its visual and olfactory senses are very limited in this habitat, and thus the possibility of predation and other risks are high in this case. Nevertheless, acoustic behavior in lesser sirens is still infrequent, and becomes even less frequent with undisturbed habitat residency, perhaps because of their familiarity with the environment. [17]

Siren intermedia make and responds to underwater sounds that may have specific communicational significance. Resting sirens are usually silent, but they may produce trains of pulsed sounds or “clicks”.  Their tendency to click is greater in groups than in single specimens, which suggests that they are involved in intraspecific communication. These acoustic behaviors occur most often when other sirens are present. They are emitted at different pulse rates by specimens clicking simultaneously, and are associated with head-jerking motions. Head-jerking may offer visual reinforcement of an acoustically defended individual space, or it may be required for sound production. S. intermedia can produce clicks by moving the horny jaw coverings together rapidly, and head-jerking may be associated with such movement when it is particularly forceful. This means of sound production is similar to the upper and lower teeth of humans, clicking with the mouth open. [17]

Another frequently produced sound ("yelp") is associated with cases of distress, or alarm. When S. intermedia butt or bite each other in their habitat, the bitten or injured individuals may swim away quickly, emitting yelps at frequencies of 880 Hz. These yelp sounds could have communicational significance if they prevent further attack or signal to other individuals that one is injured. [17]

Osmoregulation

Animals that inhabit freshwater habitats have high-affinity sodium uptake systems. Therefore, the better an animal is adapted to freshwater, the lower the rate of sodium loss and uptake. Sirens have a high sodium affinity (around 0.2 peq/g per day), and thus are able to resist the harmful effects of low-sodium environments. [8]

Hormones

Insulin has been isolated from the pancreas of lesser sirens that is mostly conserved in sequence as compared to other amphibian orders. However, it does have some substitution differences that indicate that sirens diverged early from other salamanders. [18]

Diet

Lesser sirens are filter feeders. They sift through pond bottoms and aquatic vegetation for prey items. [13] They have heavily keratinized beaks and vomerine dental arrays that can inflict substantial wounds. [7] The diet of the lesser siren includes at least 10 different taxa, which includes primarily aquatic invertebrates, including tadpoles and snails. They also eat several insects and their larvae, crayfish, mollusks, amphibians, siren eggs, and algae, although the plant material may be an incidental result of their gape-and-suck feeding style. [19]

Reproduction and life cycle

Annual seasonal behavior changes

The lesser siren is nocturnal, spending its days hidden in the debris and mud at the bottom of slow-moving bodies of water. Sirens are most active in the fall and spring seasons, when water temperatures are close to 15 °C and there is no danger of drought. [20] During the summer months, the ponds and other habitats that the lesser siren live in go through periods of drought, which leads to low water levels and possibly dried-up ponds. The lesser siren’s strategy to withstand the rigors of these dry seasons is something called “aestivation”, or summer dormancy, which can last up to 35 weeks, depending on the severity of the drought. During the sixteen-week period from July to October, aquatic lesser sirens increase the osmotic concentration of their body fluids, and slow down bodily functions. [21]

They do this by burrowing themselves into the bottom of its drying pond in tube-like channels about the length and width of their bodies. For the rest of the aestivating process, they then secrete a mucous cocoon. That is then followed by a significant drop in their oxygen consumption and heart rate. The gills slowly atrophy and the body shrinks, and as a result the fat is metabolized at one-fifth the normal rate. As expected, large individuals store more fat and consume less oxygen per unit weight than small ones, and thus can survive much longer periods of aestivation. Some individuals, especially the smaller sirens during aestivation, die or suffer from dehydration as they are unable to store sufficient fat and the greater metabolic demand reduces their chance of survival. [21]

As the drought season comes to an end, lesser sirens become active within a day, and slowly regain the lost weight over the next 8–11 weeks. [21]

Mating

Mate choice

From November to January, males occupy a shelter as a nest site usually surrounded by vegetation or among plant roots, and actively equip it with moss from the surrounding nest site, plucking plant material and dragging it back to the nest site. They then defend their territory aggressively against others through biting. [4]

Courting

Though little is known about their courtship, it is believed to be quite violent, as many specimens collected have scarring from healed bite marks from other sirens. About 12-300 eggs are laid at a time, and several clutches may be laid over the course of the year. Hatchlings are only about 0.4 in (1.1 cm) in length, but grow quickly. Maturity is typically reached in approximately three to four years. Courtship includes several repeating behavioral patterns. Once the female approaches the nest site, the male and female start moving within the area, coiling around each other. The male pursues the female’s cloaca closely, sometimes rubbing his head against the flank and the cloacal region of the female. Both the male and the female wave their tail fins by undulating the tail tip. [4]

Parental care

Oviposition

During oviposition, the female turns on its back, positions the cloaca near the top of the nest cavity and halts for several seconds. Then the male positions his cloaca near the site of oviposition. Eggs are directly coiled into the moss that the male has prepared in the nest. Interlacing of the eggs into the plant material in the nest ensures that the eggs adhere to the nest as a compact mass, and facilitates external fertilization. [4]

Egg guarding

In S. intermedia, parental care is fulfilled by the males. Paternal care as observed in the lesser siren is very rare for salamanders. After the female completes oviposition and leaves the nest, the male stays in close proximity to the eggs and takes the responsibility of parental care. The total number of eggs can be around 120–130, with a diameter of about 3 mm. The male constantly moves the egg mass, circles around it, and aerates the eggs through vigorous tail fanning. The tail-fanning behavior of the male towards the eggs can enhance sperm dispersal. Additionally, the male continuously cleans the nest from sand and other materials, to improve hygienic conditions and possibly to prevent infections from pathogens and fungi. It is also likely that males remove dead or infected eggs, in order to prevent further infection of the viable eggs. [4]

The development of the larvae takes approximately 35 days. Paternal care does not end there however, it continues after the larvae hatch and the male continues to aggressively defend the larvae up to one week after hatching. [4]

Mutualisms

Siren intermedia has many important effects on community structure as it is one of the most important predators in temporary ponds, where it complements the keystone predator role of eastern newts ( Notophthalmus viridescens ). [22] S. intermedia reduces the total densities of the anuran larvae, and by doing that, it allows the eastern newts to act as keystone predators over a broad range of prey densities. [7]

Enemies

Adult mole salamanders ( Ambystoma talpoideum ) and lesser sirens (Siren intermedia), are the top two predators in temporary ponds of the southeastern United States. Siren intermedia competes with and is an intraguild predator of A. talpoideum, limiting its growth and controlling its recruitment. [7]

Related Research Articles

<span class="mw-page-title-main">Amphibian</span> Class of ectothermic tetrapods

Amphibians are ectothermic, anamniotic, four-limbed vertebrate animals that constitute the class Amphibia. In its broad sense, it is paraphyletic group encompassing all tetrapods excluding the amniotes. All extant (living) amphibians belong to the monophyletic subclass Lissamphibia, with three living orders: Anura (frogs), Urodela (salamanders), and Gymnophiona (caecilians). Evolved to be mostly semiaquatic, amphibians have adapted to inhabit a wide variety of habitats, with most species living in freshwater, wetland or terrestrial ecosystems. Their life cycle typically starts out as aquatic larvae with gills known as tadpoles, but some species have developed behavioural adaptations to bypass this.

<span class="mw-page-title-main">Salamander</span> Order of amphibians

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 extant salamander families are grouped together under the order Urodela from the group Caudata. Salamander diversity is highest in eastern North America, especially in the Appalachian Mountains; most species are found in the Holarctic realm, with some species present in the Neotropical realm.

<i>Amphiuma</i> Genus of amphibians

Amphiuma is a genus of aquatic salamanders from the United States, the only extant genus within the family Amphiumidae. They are colloquially known as amphiumas. They are also known to fishermen as "conger eels" or "Congo snakes", which are zoologically incorrect designations or misnomers, since amphiumas are actually salamanders, and not fish, nor reptiles and are not from Congo. Amphiuma exhibits one of the largest complements of DNA in the living world, around 25 times more than a human.

<span class="mw-page-title-main">Spotted salamander</span> Species of amphibian

The spotted salamander or yellow-spotted salamander is a mole salamander common in eastern United States and Canada. It is the state amphibian of Ohio and South Carolina. The species ranges from Nova Scotia, to Lake Superior, to southern Georgia and Texas. Its embryos have been found to have symbiotic algae living in and around them, the only known example of vertebrate cells hosting an endosymbiont microbe.

<span class="mw-page-title-main">Sirenidae</span> Family of amphibians

Sirenidae, the sirens, are a family of neotenic aquatic salamanders. Family members have very small fore limbs and lack hind limbs altogether. In one species, the skeleton in their fore limbs is made of only cartilage. In contrast to most other salamanders, they have external gills bunched together on the neck in both larval and adult states. Sirens are found only in the Southeastern United States and northern Mexico.

<span class="mw-page-title-main">Greater siren</span> Species of amphibian

The greater siren is an amphibian and one of the three members of the genus Siren. The largest of the sirens and one of the largest amphibians in North America, the greater siren resides in the coastal plains of the southeastern United States.

<span class="mw-page-title-main">Two-toed amphiuma</span> Species of amphibian

The two-toed amphiuma is an aquatic salamander widely distributed in the southeastern United States. It is commonly, but incorrectly, called "congo snake", "conger eel" or the "blind eel".

<span class="mw-page-title-main">Eastern newt</span> Species of amphibian

The eastern newt is a common newt of eastern North America. It frequents small lakes, ponds, and streams or nearby wet forests. The eastern newt produces tetrodotoxin, which makes the species unpalatable to predatory fish and crayfish. It has a lifespan of 12 to 15 years in the wild, and it may grow to 5 in (13 cm) in length. These animals are common aquarium pets, being either collected from the wild or sold commercially. The striking bright orange juvenile stage, which is land-dwelling, is known as a red eft. Some sources blend the general name of the species and that of the red-spotted newt subspecies into the eastern red-spotted newt.

<span class="mw-page-title-main">Ringed salamander</span> Species of amphibian

The ringed salamander is a species of mole salamander native to hardwood and mixed hardwood-pine forested areas in and around the Ozark Plateau and Ouachita Mountains of Arkansas, Oklahoma, and Missouri. This species of salamander has slander body, small head, and long tail. They are usually found to have various dorsal color from dark gray to dark brown. Various close relatives are found such as marbled salamander and spotted salamander. This species of salamander has cannibal behavior especially those in large body size.

<span class="mw-page-title-main">Four-toed salamander</span> Species of amphibian

The four-toed salamander is a lungless salamander native to eastern North America. It is the only species of the monotypic genus Hemidactylium.

<span class="mw-page-title-main">Marbled salamander</span> Species of amphibian

The marbled salamander is a species of mole salamander found in the eastern United States.

<i>Siren</i> (genus) Genus of amphibians

Siren is a genus of aquatic salamanders of the family Sirenidae. The genus consists of five living species, along with one extinct species from the Eocene Epoch and three from the Miocene.

<span class="mw-page-title-main">San Marcos salamander</span> Species of amphibian

The San Marcos salamander is a small species of aquatic, lungless salamander native to the United States, endemic to Spring Lake and a small region of the headwaters of the San Marcos River near Aquarena Springs, in Hays County, Texas. It is one to two inches long, with a slender body and external gills, and is reddish-brown in color.

<i>Ambystoma talpoideum</i> Species of salamander

Ambystoma talpoideum, the mole salamander, is a species of salamander found in much of the eastern and central United States, from Florida to Texas, north to Illinois, east to Kentucky, with isolated populations in Virginia and Indiana. Older sources often refer to this species as the tadpole salamander because some individuals remain in a neotenic state. This salamander lives among the leaf litter on the forest floor, migrating to ponds to breed.

<span class="mw-page-title-main">Southern dwarf siren</span> Species of amphibian

The southern dwarf siren, is a perennibranch salamander lacking hind legs. Found exclusively in Florida, it is one of two currently recognized species of dwarf sirens. Two subspecies are currently recognized; P. a. axanthus, the narrow-striped dwarf siren, and P. a. belli, the Everglades dwarf siren.

<span class="mw-page-title-main">Cheat Mountain salamander</span> Species of amphibian

The Cheat Mountain salamander is a species of small woodland salamander in the family Plethodontidae. The species is found only on Cheat Mountain, and a few nearby mountains, in the eastern highlands of West Virginia. It and the West Virginia spring salamander are the only vertebrate species with geographic ranges restricted to that state.

<span class="mw-page-title-main">Oita salamander</span> Species of amphibian

The Oita salamander is a species of salamander in the family Hynobiidae endemic to Japan. Named after Ōita Prefecture, its natural habitats are temperate forests, rivers, intermittent rivers, freshwater marshes, intermittent freshwater marshes, and irrigated land in western Japan. It is threatened by habitat loss, due to the increasing construction of homes within its habitat. The Oita salamander is considered to be vulnerable by the (IUCN) Red List of Threatened Species with a declining population.

<i>Desmognathus fuscus</i> Species of amphibian

Desmognathus fuscus is a species of amphibian in the family Plethodontidae. 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. The northern dusky salamander is the most widespread representative of its genus in Canada. It can be found in eastern North America from extreme eastern Canada in New Brunswick south to South Carolina. 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, 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 larval stage which follows is normally aquatic.

<span class="mw-page-title-main">Newt</span> Salamander in the subfamily Pleurodelinae

A newt is a salamander in the subfamily Pleurodelinae. The terrestrial juvenile phase is called an eft. Unlike other members of the family Salamandridae, newts are semiaquatic, alternating between aquatic and terrestrial habitats. Not all aquatic salamanders are considered newts, however. More than 100 known species of newts are found in North America, Europe, North Africa and Asia. Newts metamorphose through three distinct developmental life stages: aquatic larva, terrestrial juvenile (eft), and adult. Adult newts have lizard-like bodies and return to the water every year to breed, otherwise living in humid, cover-rich land habitats.

<i>Speleomantes strinatii</i> Species of amphibian

Speleomantes strinatii, the French cave salamander, North-west Italian cave salamander, or Strinati's cave salamander is a small species of salamander found in northwest Italy and southeast France. It is very similar in appearance to the Italian cave salamander, but has a paler belly.

References

  1. 1 2 3 Parra-Olea, G.; Wake, D.; Hammerson, G.A. (2008). "Siren intermedia". IUCN Red List of Threatened Species . 2008: e.T59491A11936674. doi: 10.2305/IUCN.UK.2008.RLTS.T59491A11936674.en . Retrieved 19 November 2021.
  2. ↑ Lesser siren (siren intermedia). Species Profile: Lesser Siren (Siren intermedia) | SREL Herpetology. (n.d.). Retrieved April 28, 2022, from https://srelherp.uga.edu/salamanders/sirint.htm
  3. 1 2 Caudata Culture Species Entry – Siren intermedia. Caudata.org. Retrieved on 2019-12-23.
  4. 1 2 3 4 5 6 Reinhard, Sandy, et al. “External Fertilisation and Paternal Care in the Paedomorphic Salamander Siren Intermedia Barnes, 1826 (Urodela: Sirenidae).” Zoologischer Anzeiger - a Journal of Comparative Zoology, vol. 253, no. 1, August 2013, pp. 1–5, 10.1016/j.jcz.2013.06.002. Accessed 25 March 2022.
  5. ↑ Graham, Sean P.; Kline, Richard; Steen, David A.; Kelehear, Crystal (5 December 2018). "Description of an extant salamander from the Gulf Coastal Plain of North America: The Reticulated Siren, Siren reticulata". PLOS ONE. 13 (12): e0207460. Bibcode:2018PLoSO..1307460G. doi: 10.1371/journal.pone.0207460 . ISSN   1932-6203. PMC   6281224 . PMID   30517124.
  6. ↑ Tipton, Bob L. (2012). Texas amphibians : a field guide (1st ed.). Austin. ISBN   978-0292737358.{{cite book}}: CS1 maint: location missing publisher (link)
  7. 1 2 3 4 Fauth, John E. “Interactions between Branchiate Mole Salamanders (Ambystoma Talpoideum) and Lesser Sirens (Siren Intermedia): Asymmetrical Competition and Intraguild Predation.” Amphibia-Reptilia, vol. 20, no. 2, 1 May 1999, pp. 119–132, 10.1163/156853899506942. Accessed 30 October 2020.
  8. 1 2 3 Asquith, Adam, and Ronald Altig. “Osmoregulation of the Lesser Siren, Siren Intermedia (Caudata: Amphibia).” Comparative Biochemistry and Physiology Part A: Physiology, vol. 84, no. 4, January 1986, pp. 683–685, 10.1016/0300-9629(86)90387-7. Accessed 1 May 2021.
  9. ↑ "Siren intermedia". amphibiaweb.org. AmphibiaWeb. Retrieved 26 December 2018.
  10. ↑ "Siren intermedia". amphibiaweb.org. AmphibiaWeb. Retrieved 17 December 2018.
  11. 1 2 "Siren intermedia nettingi (Western lesser siren)". Michigan Herp Atlas. Herpetological Resource and Management / Michigan DNR. Retrieved 4 August 2019.
  12. ↑ Williams, J. E. (2004). A first record of the Western Lesser Siren, Siren intermedia nettingi, from Michigan. (Fisheries research report: 1588).
  13. 1 2 3 Anderson, Thomas L., et al. “Functional Responses of Larval Marbled Salamanders (Ambystoma Opacum) and Adult Lesser Sirens (Siren Intermedia) on Anuran Tadpole Prey.” Copeia, vol. 108, no. 2, 19 May 2020, p. 341, 10.1643/ce-19-212. Accessed 20 February 2022.
  14. ↑ Gehlbach, Frederick R., and Stephen E. Kennedy. “Population Ecology of a Highly Productive Aquatic Salamander (Siren Intermedia).” The Southwestern Naturalist, vol. 23, no. 3, 10 August 1978, p. 423, 10.2307/3670250. Accessed 14 May 2021.
  15. ↑ "Western Lesser Siren (Siren intermedia nettingi)". www.miherpatlas.org. Retrieved 2019-08-04.
  16. ↑ • MARTIN, B. T., GOODDING, D. D., FORD, N. B., & PLACYK, J. . J. S. (2013). Sensory Mediation of Foraging Behavior in the Western Lesser Siren (Siren intermedia nettingi). Journal of Herpetology, 47(1), 75–77. https://doi.org/10.1670/11-209
  17. 1 2 3 Gehlbach, Frederick R., and Braz Walker. “Acoustic Behavior of the Aquatic Salamander, Siren Intermedia.” BioScience, vol. 20, no. 20, 15 October 1970, pp. 1107–1108, 10.2307/1295417. Accessed 11 March 2021.
  18. ↑ Conlon, J. Michael; Trauth, Stanley E.; Sever, David M. (1997-06-01). "Purification and Structural Characterization of Insulin from the Lesser Siren,Siren intermedia(Amphibia: Caudata)". General and Comparative Endocrinology. 106 (3): 295–300. doi:10.1006/gcen.1997.6912. ISSN   0016-6480. PMID   9204362.
  19. ↑ Hampton, Paul M. “Ecology of the Lesser Siren, Siren Intermedia, in an Isolated Eastern Texas Pond.” Journal of Herpetology, vol. 43, no. 4, December 2009, pp. 704–709, 10.1670/08-168.1. Accessed 25 March 2022.
  20. ↑ Martin, Bradley T., et al. “Sensory Mediation of Foraging Behavior in the Western Lesser Siren (Siren Intermedia Nettingi).” Journal of Herpetology, vol. 47, no. 1, March 2013, pp. 75–77, 10.1670/11-209. Accessed 25 March 2022.
  21. 1 2 3 Gehlbach, Frederick R., et al. “Aestivation of the Salamander, Siren Intermedia.” American Midland Naturalist, vol. 89, no. 2, April 1973, p. 455, 10.2307/2424051. Accessed 25 March 2022.
  22. ↑ Fauth, John E.; Resetarits, William J. (June 1991). "Interactions Between the Salamander Siren Intermedia and the Keystone Predator Notophthalmus Viridescens". Ecology. 72 (3): 827–838. doi:10.2307/1940585. JSTOR   1940585.
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