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Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Cladus: Craniata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Lepidosauromorpha
Superordo: Lepidosauria
Ordo: Squamata
Subordo: Serpentes
Infraordo: Caenophidia
Superfamilia: Viperoidea

Familia: Viperidae
Subfamilia: Causinae
Genus: Causus
Species: C. bilineatus - C. defilippii - C. lichtensteinii - C. maculatus - C. resimus - C. rhombeatus

Name

Causus Wagler, 1830

Type species: Causus rhombeatus (Lichtenstein, 1823)

Synonyms

Distichurus Hallowell, 1842
Heterophis W. Peters, 1862
Dinodipsas W. Peters, 1882

Vernacular names
Deutsch: Krötenvipern
English: Night Adders
eesti: Kärnrästik
עברית: צפע לילי
Nederlands: Padadders
norsk: Nattvipere
русский: Жабьи гадюки
українська: Жаб'яча гадюка
中文: 夜蝰屬

Causus is a genus of venomous vipers found only in sub-Saharan Africa. It is a group considered to be among the most primitive members of the family Viperidae based on head scalation, oviparity, venom apparatus, and because they have round pupils.[2][3] However, this is contradicted by recent molecular studies.[4][5] Seven species are currently recognized.[6][7] They are commonly known as night adders.[2][3]

Description

These snakes are fairly stout, never growing to more than 1 m (3.3 ft) in total length.[3]

As opposed to most vipers, where the head is distinct from the neck and covered with small scales, in Causus the head is only slightly distinct from the neck and covered with 9 large, symmetrical head shields. Also, the eyes have round pupils instead of elliptical ones like other vipers. The rostral scale is broad, sometimes pointed or upturned. The nostril is located between two nasals and an internasal. The frontal and supraocular scales are long. A loreal scale is present, separating the nasal and preoculars. The suboculars are separated from the supralabials. The mandible has splenial and angular elements.[2]

The fangs are different, too. Unlike other vipers, no hinge action occurs where the prefrontal bone engages the frontal. However, since the maxillary bones rotate almost as far, the fangs can still be erected. The fangs themselves are relatively short. A fine line, or suture, is also present along the length of the fang, representing the vestigial edge where the groove lips meet (from incomplete fang canal closure).[2]

The body is cylindrical or slightly depressed and moderately slender. The dorsal scales are smooth or weakly keeled with apical pits. The ventral scales are rounded, and the anal plate is single. The tail is short, and the subcaudals can be either single or paired.[2]

Among the viperids, another unique characteristic of this genus is several species have venom glands that are not confined to the temporal area as with most vipers, but are exceptionally long and extend well down the neck.[3] These venom glands, located on either side of the spine, may be up to 10 cm in length, with long ducts connecting them to the fangs.[8]

Other internal differences also set the Causinae apart: they have unusually long kidneys, a well-developed tracheal lung with two tracheal arteries, and the liver overlaps the tip of the heart.[2]
Geographic range
A young night adder, photographed in Kei Mouth, South Africa

The Causinae are found in sub-Saharan Africa.[1]
Behavior

Despite their common name, this genus is active during the day, as well as at night. When disturbed, they will engage in a ferocious hissing and puffing threat display. They may lift the first part of their bodies off the ground in a coil and make a powerful swiping strike; juveniles have been known to come off the ground. Others specimens may raise the first part of their bodies off the ground, flattening their necks and moving forward, tongue extended, like a small cobra.[3] The rather frantic strikes are often combined with attempts to quickly glide away.[9]
Feeding

Their diet consists almost exclusively of toads and frogs.[3] Gluttony has been reported; when prey is abundant, they may eat until they are literally unable to swallow any more food.[9]
Reproduction

All Causus species lay eggs (oviparous), which among vipers is considered to be a more primitive trait, though not unique.[2] The average clutch consists of some two dozen eggs that require an incubation period of about four months. Hatchlings are 10–12.5 cm (4–5 in) in total length (body + tail).[9]
Venom

In spite of their enormously developed venom glands, night adders do not always use them to subdue their prey. The venom would act fast enough, but often they simply seize and swallow their prey, instead.[8]

Causus venom is weaker than that of puff adder. Envenomation normally causes only local pain and swelling.[2] Antivenin treatment should not be necessary. Nevertheless, South African polyvalent serum is known to be effective against the venom of at least two species.[10] However, envenomation can be very serious and in at least one case a bitten child had to have a fasciotomy. A number of small dogs have had limbs amputated or died due to bites from this snake. The danger from this snake should not be underestimated as a bite from a large individual on a small child could potentially be fatal.[11]

No recent deaths have been reported due to this species. Earlier reports of fatalities were based on anecdotal evidence; the species involved likely were not properly identified or the cases were grossly mismanaged.[10]
Species

Species[6][7] Taxon author[7] Common name Geographic range
Causus bilineatus Boulenger, 1905 Two-striped night adder Africa in Angola, Zambia, and southern DR Congo
Causus defilippii (Jan, 1863) Snouted night adder Southeast Africa
Causus lichtensteinii (Jan, 1859) Lichtenstein's night adder Equatorial Africa from Zambia and Cameroon to the Ivory Coast
Causus maculatus (Hallowell, 1842) West African night adder Most of sub-Saharan Africa north of the equator
Causus rasmusseni Broadley, 2014 Angolan night adder Zambia and Angola
Causus resimus (W. Peters, 1862) Green night adder Most of equatorial Africa
Causus rhombeatusT (Lichtenstein, 1823) Common night adder Sub-Saharan Africa

T) Type species.
Taxonomy

A long-standing tendency among herpetologists was to make assumptions regarding the phylogenetic position of this group. McDiarmid et al.[1] suggested the consensus among the experts in 1999 was that Causus was basal to all viperids, so deserved a separate subfamily. However, two more recent molecular phylogenetic studies[4][5] have shown Causus is not only not basal to all viperids, but in fact appears to be nested within the viperines. Consequently, recognition of the subfamily Causinae is inappropriate.
References

McDiarmid RW, Campbell JA, Touré T. 1999. Snake Species of the World: A Taxonomic and Geographic Reference, Volume 1. Herpetologists' League. 511 pp. ISBN 1-893777-00-6 (series). ISBN 1-893777-01-4 (volume).
Mallow D, Ludwig D, Nilson G. 2003. True Vipers: Natural History and Toxinology of Old World Vipers. Krieger Publishing Company. Malabar, Florida. 359 pp. ISBN 0-89464-877-2.
Spawls S, Branch B. 1995. The Dangerous Snakes of Africa. Ralph Curtis Books. Dubai: Oriental Press. 192 pp. ISBN 0-88359-029-8.
Nagy ZT, Vidal N, Vences M, Branch WR, Pauwels OSG, Wink M, Joger U. 2005. Molecular systematics of African Colubroidea (Squamata: Serpentes) In: Huber BA, Sinclair BJ, Lampe KH (Eds.). African Biodiversity: Molecules, Organisms, Ecosystems. Proc. 5th Intern. Symp. Trop. Biol. Museum Koenig, Bonn, pp. 221-228.
Wüster W, Peppin L, Pook CE, Walker DE. 2008. A nesting of vipers: phylogeny and historical biogeography of the Viperidae (Squamata: Serpentes). Molecular Phylogenetics and Evolution 49: 445-459.
Causus at the Reptarium.cz Reptile Database. Accessed 20 June 2021.
"Causus". Integrated Taxonomic Information System. Retrieved 20 June 2021.
Stidworthy J. 1974. Snakes of the World. Grosset & Dunlap Inc. 160 pp. ISBN 0-448-11856-4.
Mehrtens JM. 1987. Living Snakes of the World in Color. New York: Sterling Publishers. 480 pp. ISBN 0-8069-6460-X.
Spawls S, Howell K, Drewes R, Ashe J. 2004. A Field Guide To The Reptiles Of East Africa. A & C Black Publishers Ltd. London. 543 pp. ISBN 0-7136-6817-2.

"African Snakebite Institute _ About Johan Marais". africansnakebiteinstitute.com. Archived from the original on 2017-02-09. Retrieved 2017-01-15.

Further reading

Cope ED. 1860 (dated 1859). Catalogue of the Venomous Serpents in the Museum of the Academy of Natural Sciences of Philadelphia, with notes on the Families, Genera and Species. Proc. Acad. Nat. Sci. Philadelphia 11: 332-347.
Hallowell E. 1842. Description of a new Genus of Serpents from Western Africa. Journal of the Academy of Natural Sciences of Philadelphia 8: 336-338. (Genus Distichurus)
Peters W. 1862. Über die von dem so früh in Afrika verstorbenen Freiherrn von Barnim und Dr. Hartmann auf ihrer Reise durch Aegypten, Nubien und dem Sennâr gesammelten Amphibien. Monatsberichte der Preussischen Akademie Wissenschaften zu Berlin 1862: 271-279. ("Heterophis gen. nov.", pp. 276–277.)
Peters W. 1882. Über eine neue Gattung und Art der Vipernattern, Dinodipsas angulifera , aus Südamerica. Sitzungsberichte der Preussischen Akademie Wissenschaften zu Berlin 1882 (40): 893-896. (Genus Dinodipsas, p. 893.)
Wagler J. 1830. Natürliches System der Amphibien, mit vorangehender Classification der Säugthiere und Vögel. Ein Beitrag zur vergleichenden Zoologie. J.G. Cotta. Munich, Stuttgart, and Tübingen. vi + 354 pp. + 9 plates. (Genus Causus, pp. 172–173.)

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