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Sirenidae, the sirens, are a
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of neotenic aquatic
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All t ...
s. Family members have very small fore limbs, and lack hind limbs altogether. In one species, the
skeleton A skeleton is the structural frame that supports the body of most animals. There are several types of skeletons, including the exoskeleton, which is a rigid outer shell that holds up an organism's shape; the endoskeleton, a rigid internal fra ...
in their fore limbs is made of only
cartilage Cartilage is a resilient and smooth type of connective tissue. Semi-transparent and non-porous, it is usually covered by a tough and fibrous membrane called perichondrium. In tetrapods, it covers and protects the ends of long bones at the joints ...
. In contrast to most other salamanders, they have external
gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow r ...
s bunched together on the
neck The neck is the part of the body in many vertebrates that connects the head to the torso. It supports the weight of the head and protects the nerves that transmit sensory and motor information between the brain and the rest of the body. Addition ...
in both
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
l and adult states. Sirens are found only in the
Southeastern United States The Southeastern United States, also known as the American Southeast or simply the Southeast, is a geographical List of regions in the United States, region of the United States located in the eastern portion of the Southern United States and t ...
and northern
Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...
. Although they are primarily
carnivorous A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose nutrition and energy requirements are met by consumption of animal tissues (mainly mu ...
, they are the only salamanders observed eating plant material.Hill, R. L., Mendelson, J. R. & Stabile, J. L. 2015. Direct observation and review of herbivory in Sirenidae (Amphibia: Caudata). Southeastern Naturalist 14, N5-N9.


Description

Sirens are quite distinct from other salamanders, and in some classifications they form their own suborder, Sirenoidea, or as a completely distinct order (Meantes or Trachystomata). Genetic analysis variously places them as the sister to other
Salamandroidea The Salamandroidea are a suborder of salamanders, referred to as advanced salamanders. The members of the suborder are found worldwide except for Antarctica, sub-Saharan Africa, and Oceania. They differ from suborder Cryptobranchoidea as the Angu ...
or as sister to all other salamanders. Many of their unique characteristics seem to be partly primitive and partly derivative. Sirens are generally eel-like in form, with two tiny, but otherwise fully developed, fore limbs. They range from in length. They are neotenic, although the larval
gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow r ...
s are small and functionless at first, and only adults have fully developed (but inefficient) gills. They are obligate air-breathers with well developed lungs. Proving they likely evolved from a terrestrial ancestor with an aquatic larval stage. Like
amphiuma ''Amphiuma'' is a genus of aquatic salamanders from the United States, the only Extant taxon, extant genus within the family (biology), family Amphiumidae . They are colloquially known as amphiumas. They are also known to fishermen as "conger eel ...
s, they are able to cross land on rainy nights. These amphibians are
omnivorous An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize ...
, feeding mainly on
worm Worms are many different distantly related bilateria, bilateral animals that typically have a long cylindrical tube-like body, no limb (anatomy), limbs, and usually no eyes. Worms vary in size from microscopic to over in length for marine ...
s, small
snail A snail is a shelled gastropod. The name is most often applied to land snails, terrestrial molluscs, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gas ...
s,
shrimp A shrimp (: shrimp (American English, US) or shrimps (British English, UK)) is a crustacean with an elongated body and a primarily Aquatic locomotion, swimming mode of locomotion – typically Decapods belonging to the Caridea or Dendrobranchi ...
s, and filamentous
algae Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthesis, photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular ...
.John Farrand Jr., The Audubon Society Encyclopedia of Animal Life, 1982 They are notable among salamanders (and most amphibians, aside from a few frog species) due to their semi-
herbivorous A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat n ...
habits. Their jaws possess sharp-edged keratinised and toothless ridges, like many anuran tadpoles, but the coronoid bone in the lower jaw and the
vomer The vomer (; ) is one of the unpaired facial bones of the skull. It is located in the midsagittal line, and articulates with the sphenoid, the ethmoid, the left and right palatine bones, and the left and right maxillary bones. The vomer forms ...
and
palatine bone In anatomy, the palatine bones (; derived from the Latin ''palatum'') are two irregular bones of the facial skeleton in many animal species, located above the uvula in the throat. Together with the maxilla, they comprise the hard palate. Stru ...
in the upper jaw have patches of monocuspid and unbladed teeth arranged in polystichous patterns. If the conditions of a water source are unsuitable, a larva will shrink its gills to mere stumps, and these may not function at all. They are also able to burrow into mud of drying
pond A pond is a small, still, land-based body of water formed by pooling inside a depression (geology), depression, either naturally or artificiality, artificially. A pond is smaller than a lake and there are no official criteria distinguishing ...
s and encase themselves with a cocoon of
mucus Mucus (, ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both Serous fluid, serous and muc ...
to survive periods of drought. During such periods, they breathe with their small but functional
lung The lungs are the primary Organ (biology), organs of the respiratory system in many animals, including humans. In mammals and most other tetrapods, two lungs are located near the Vertebral column, backbone on either side of the heart. Their ...
s. Unlike other salamanders, an
interventricular septum The interventricular septum (IVS, or ventricular septum, or during development septum inferius) is the stout wall separating the ventricle (heart), ventricles, the lower chambers of the heart, from one another. The interventricular septum is di ...
is present in the heart. At least two of the species can produce vocalizations. The structure of sirens' reproductive systems suggests they employ external fertilization. This has finally been confirmed in captive breeding experiments, showcasing that males also engage in
parental care Parental care is a behavioural and evolutionary strategy adopted by some animals, involving a parental investment being made to the evolutionary fitness of offspring. Patterns of parental care are widespread and highly diverse across the animal k ...
, building nests for their offspring.Reinhard, S., Voitel, S. & Kupfer, A. 2013. External fertilisation and paternal care in the paedomorphic salamander Siren intermedia Barnes, 1826 (Urodela: Sirenidae). Zoologischer Anzeiger 253, 1–5. Parental care among sirens is paternal due to external fertilization. In ''S. intermedia'' males circle around females and may rub or bite her flank region. Both male and female will go on their backs and turn. It is assumed here where the female spawns and the male fertilizes her eggs. After the courtship is over, the female leaves and the male guards the eggs. Males could potentially guard more than one brood, but they are known to bite females who enter a nesting site. Paternal care has also been observed in
Cryptobranchoidea The Cryptobranchoidea are a suborder of salamanders found in Asia, European Russia, and the United States. They are known as primitive salamanders, in contrast to Salamandroidea, the advanced salamanders. It has two living subdivisions, Cryptobr ...
, the other suborder with external fertilization. This is critical to phylogeny, as
Salamandroidea The Salamandroidea are a suborder of salamanders, referred to as advanced salamanders. The members of the suborder are found worldwide except for Antarctica, sub-Saharan Africa, and Oceania. They differ from suborder Cryptobranchoidea as the Angu ...
, the third suborder, use internal fertilization which may be pair with maternal care, meaning that sirens are one of the oldest groups of salamanders. The combined
biomass Biomass is a term used in several contexts: in the context of ecology it means living organisms, and in the context of bioenergy it means matter from recently living (but now dead) organisms. In the latter context, there are variations in how ...
of ''
Siren intermedia 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 ep ...
'' species in a Texas pond exceeded the total biomass of the pond's seven species of fish.


Taxonomy

The siren family (Sirenidae) is subdivided into five genera, three extinct, and two extant with two and three extant species, respectively: * Genus †'' Habrosaurus'' Gilmore 1928 ** †'' H. dilatus'' Gilmore 1928 ** †'' H. prodilatus'' Gardner 2003 * Genus †'' Kababisha'' Evans et al. 1996 ** †'' K. humarensis'' Evans et al. 1996 ** †'' K. sudanensis'' Evans et al. 1996 * Genus †'' Noterpeton'' Rage et al. 1993 ** †'' Noterpeton bolivianum'' Rage et al. 1993 * Genus '' Pseudobranchus'' Gray 1825 dwarf sirens ** †'' P. robustus'' Goin and Auffenberg 1955 ** †'' P. vetustus'' ** '' P. axanthus''
Netting In law, set-off or netting is a legal technique applied between persons or businesses with mutual rights and Liability (financial accounting), liabilities, replacing gross positions with net positions. It permits the rights to be used to discharg ...
& Goin 1942
southern dwarf siren ** '' P. striatus'' LeConte 1824 northern dwarf siren * Genus '' Siren'' Österdam 1766 sirens ** †'' S. dunni'' Goin and Auffenberg 1957 ** †'' S. hesterna'' ** †'' S. miotexana'' ** †'' S. simpsoni'' ** '' S. intermedia'' Barnes 1826 lesser siren ** '' S. lacertina''
Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
, 1766
greater siren ** ''S. nettingi'' Goin, 1942 western siren **'' S. reticulata'' Graham, Kline, Steen & Kelehear, 2018 reticulated siren **''S. sphagnicola'' Fedler, Enge, & Moler, 2023 seepage siren


References

*


External links


Tree of Life: Sirenidae
* {{Authority control Cenomanian first appearances Taxa named by John Edward Gray Amphibian families Extant Cenomanian first appearances