Atheris

{{Short description|Genus of venomous vipers of tropical Africa}} {{Update|inaccurate=yes|date=June 2011}} {{Automatic taxobox | fossil_range = Miocene–present {{Fossil range|15|0}} <ref>{{Cite journal |last1=Menegon |first1=M. |last2=Loader |first2=S. P. |last3=Marsden |first3=S. J. |last4=Branch |first4=W. R. |last5=Davenport |first5=T. R. B. |last6=Ursenbacher |first6=S. |date=2014-10-01 |title=The genus Atheris (Serpentes: Viperidae) in East Africa: Phylogeny and the role of rifting and climate in shaping the current pattern of species diversity |url=https://www.sciencedirect.com/science/article/abs/pii/S1055790314002206 |journal=Molecular Phylogenetics and Evolution |volume=79 |pages=12–22 |doi=10.1016/j.ympev.2014.06.007 |pmid=24952316 |bibcode=2014MolPE..79...12M |issn=1055-7903|url-access=subscription }}</ref> | image = Atheris hispida.jpg | image_caption = ''Atheris hispida'' | taxon = Atheris | authority = Cope, 1862 | synonyms = * ''Chloroechis'' Bonaparte, 1849 * ''Atheris'' Cope, 1862 * ''Poecilostolus'' Günther, 1863<ref name="McD99">McDiarmid RW, Campbell JA, Touré T (1999). ''Snake Species of the World: A Taxonomic and Geographic Reference, Volume 1''. Washington: District of Columbia: Herpetologists' League. 511 pp. {{ISBN|1-893777-00-6}} (series). {{ISBN|1-893777-01-4}} (volume).</ref> }}

'''''Atheris''''' is a genus of vipers known as '''bush vipers'''.<ref name="SB95">Spawls S, Branch B (1995). ''The Dangerous Snakes of Africa''. Ralph Curtis Books. Dubai: Oriental Press. 192 pp. {{ISBN|0-88359-029-8}}.</ref> They are found only in tropical subsaharan Africa (excluding southern Africa)<ref name="McD99"/> and many species have isolated and fragmented distributions due to their confinement to rain forests.<ref name="Mal03">Mallow D, Ludwig D, Nilson G (2003). ''True Vipers: Natural History and Toxinology of Old World Vipers''. Malabar, Florida: Krieger Publishing Company. 359 pp. {{ISBN|0-89464-877-2}}.</ref> Like all other vipers, they are venomous. In an example of convergent evolution, they show many similarities to the arboreal pit vipers of Asia and South America.<ref name="SB95"/> Seventeen species are currently recognized.<ref name="ITIS">{{ITIS |id=634419 |taxon=''Atheris'' |accessdate=13 July 2006}}</ref>

==Description== They are relatively small in size, with adults ranging in total length (body + tail) from {{convert|55|cm|in|abbr=on}} for ''A. katangensis'' to a maximum of {{convert|78|cm|in|abbr=on}} for ''A. squamigera''.<ref name="SB95"/>

All species have a broad, triangular head that is distinct from the neck. The canthus is also distinct and the snout is broad. The crown is covered with small imbricate or smooth scales, none of which is enlarged. The eyes are relatively large with elliptical pupils. The eyes are separated from the supralabials by 1–3 scale rows and from the nasal by 2–3 scales.<ref name="Mal03"/>

The body is slender, tapering, and slightly compressed. The dorsal scales are overlapping, strongly keeled and have apical pits. Laterally these are smaller than the middorsals. Midbody there are 14–36 rows of dorsal scales. There are 133–175 rounded ventral scales. The subcaudal scales are single and number 38–67.<ref name="SB95"/><ref name="Mal03"/> The tail is strongly prehensile and can support the body while suspended from a branch or a twig.<ref name="Meh87">Mehrtens JM (1987). ''Living Snakes of the World in Color''. New York: Sterling Publishers. 480 pp. {{ISBN|0-8069-6460-X}}.</ref>

Members of this genus come in a wide variety of colors and patterns, often within a single species. ''A. Ceratophora'' and ''A. Squamigera'' are particularly variable.<ref name="WOAO">{{WOA|page=overview|subject=Overview|date=8 September|year=2007}}</ref>

==Location== They inhabit rainforest regions and forests, mostly in remote areas far from human activity.<ref name="SB95"/>

They are found in tropical subsaharan Africa, excluding southern Africa.<ref name="McD99"/> Some species have only isolated populations, surviving in small sections of ancient rainforest. They once had a much wider distribution but are now declining.<ref name="SB95"/>

==Conservation status== Some species are threatened by habitat destruction.<ref name="SB95"/> A major cause of illness and mortality in both caged and wild bush viper snakes is snake fungal disease.<ref name="Diaz2020">{{cite journal |last1=Díaz-Delgado |first1=Josué |last2=Marrow |first2=Judilee C. |last3=Flanagan |first3=Joseph P. |last4=Bauer |first4=Kendra L. |last5=Zhang |first5=Meng |last6=Rodrigues-Hoffmann |first6=Aline |last7=Groch |first7=Katia R. |last8=Gomez |first8=Gabriel |last9=Balamayooran |first9=Gayathriy |title=Outbreak of Paranannizziopsis australasiensis Infection in Captive African Bush Vipers (Atheris squamigera) |url=https://www.sciencedirect.com/science/article/abs/pii/S0021997520301390 |journal=Journal of Comparative Pathology |access-date=20 March 2022 |pages=97–102 |language=en |doi=10.1016/j.jcpa.2020.10.004 |date=1 November 2020|volume=181 |pmid=33288159 |s2cid=227955419 |url-access=subscription }}</ref>

==Behavior== All species have extreme aggressive tendencies. All species are arboreal, although they can sometimes be found on or near the ground.<ref name="WOAO"/>

''Atheris'' species have been known to prey upon a variety of small amphibians, lizards, rodents, birds, and even other snakes. Some species or populations may specialize in eating frogs, but most have been described as opportunistic feeders.<ref name="Mal03"/><ref name="WOAO"/> Prey is typically ambushed from a hanging position, held until it has succumbed to the venom, and then swallowed.<ref name="WOAO"/>

All ''Atheris'' species are ovoviviparous.<ref name="Meh87" /> Mating takes place in October and November, and the females give birth to live young in March and April.<ref name="WOA-C">{{WOA|page=captivity|subject=Captivity|date=8 September|year=2007}}</ref>

==Captivity== ''A. squamigera'' is reported to do very well in captivity, needing only arboreal access and having no particular temperature requirements. Captive specimens take mice and small birds.<ref name="Mal03"/> However, there have been reports of cannibalism.<ref name="WOAO"/> Food may be refused during the African winter months of July and August.

==Venom== Limited understanding exists about their venom, except that it is strongly hemotoxic, causing pain, swelling, and blood clotting problems.<ref name="SB95"/> Until recently, their venom has often been regarded as less toxic than that of many other species, perhaps because bites are uncommon,<ref name="Mal03"/> but this turned out not to be the case. There are now a number of reports of bites that have led to severe hemorrhaging.<ref name="Meb98">{{cite journal |doi=10.1016/S0041-0101(98)00008-7 |vauthors=Mebs D, Holada K, Kornalík F |title=Severe coagulopathy after a bite of a green bush viper (''Atheris squamiger''): case report and biochemical analysis of the venom |journal=Toxicon |volume=36 |issue=10 |pages=1333–40 |date=October 1998 |pmid=9723832 |bibcode=1998Txcn...36.1333M |display-authors=etal}}</ref><ref name="Top06">{{cite journal|vauthors=Top LJ, Tulleken JE, ((Ligtenberg JJM)), ((Meertens JHJM)), van der Werf TS, Zijlstra JG |title=Serious envenomation after a snakebite by a Western bush viper (''Atheris chlorechis'') in the Netherlands: a case report |journal=Neth. J. Med. |volume=64 |issue=5 |pages=153–6 |year=2006 |url=http://www.njmonline.nl/njm/cntpdf.php?t=a&id=10000066 |format=PDF |pmid=16702615 }}{{dead link|date=October 2016 |bot=InternetArchiveBot |fix-attempted=yes }}</ref><ref name="VR00">[http://www.venomousreptiles.org/articles/28 Bitten by a Sedge Viper!] {{Webarchive|url=https://web.archive.org/web/20091210044333/http://www.venomousreptiles.org/articles/28 |date=2009-12-10 }} at [http://www.venomousreptiles.org/ VenomousReptiles.org] {{Webarchive|url=https://web.archive.org/web/20080409033049/http://www.venomousreptiles.org/ |date=2008-04-09 }}. Accessed 2 August 2007.</ref> A case of a bite from ''A. nitschei'' caused local tissue injury, hemolysis, thrombocytopenia, hypofibrinoginemia, coagulopathy, and increased elevation in D-dimer. <ref>{{Cite journal |last1=Hatten |first1=Benjamin W. |last2=Bueso |first2=Antonio |last3=French |first3=L. Keith |last4=Hendrickson |first4=Robert G. |last5=Horowitz |first5=B. Zane |date=2013-02-01 |title=Envenomation by the great lakes bush viper (Atheris nitschei) |url=http://www.tandfonline.com/doi/full/10.3109/15563650.2012.763134 |journal=Clinical Toxicology |language=en |volume=51 |issue=2 |pages=114–116 |doi=10.3109/15563650.2012.763134 |pmid=23327286 |issn=1556-3650|url-access=subscription }}</ref> Envenomation cases from ''A. chlorechis'' and ''A. squamigera'' have shown similar symptoms.<ref>{{Cite journal |last1=Top |first1=L. J. |last2=Tulleken |first2=J. E. |last3=Ligtenberg |first3=J. J. M. |last4=Meertens |first4=J. H. J. M. |last5=van der Werf |first5=T. S. |last6=Zijlstra |first6=J. G. |date=May 2006 |title=Serious envenomation after a snakebite by a Western bush viper (Atheris chlorechis) in the Netherlands: a case report |journal=The Netherlands Journal of Medicine |volume=64 |issue=5 |pages=153–156 |issn=0300-2977 |pmid=16702615}}</ref><ref>{{Cite journal |last1=Mebs |first1=D. |last2=Holada |first2=K. |last3=Kornali ́k |first3=F. |last4=Simák |first4=J. |last5=Vanková |first5=H. |last6=Müller |first6=D. |last7=Schoenemann |first7=H. |last8=Lange |first8=H. |last9=Herrmann |first9=H-W. |date=1998-10-01 |title=Severe coagulopathy after a bite of a green bush viper (Atheris squamiger): Case report and biochemical analysis of the venom |url=https://www.sciencedirect.com/science/article/abs/pii/S0041010198000087 |journal=Toxicon |volume=36 |issue=10 |pages=1333–1340 |doi=10.1016/S0041-0101(98)00008-7 |pmid=9723832 |bibcode=1998Txcn...36.1333M |issn=0041-0101|url-access=subscription }}</ref> One case was fatal.<ref name="Mal03"/> ''Atheris''-specific antivenom does not exist<ref name="SB95"/> and antivenom meant for bites from other species seems to have little effect, although ''Echis'' antivenom has been reported to have been of some help in a case of ''A. squamigera'' envenomation.<ref name="Mal03"/> Symptomatic treatment is all that can be administered due to the absence of an Atheris-specific antivenom.<ref>Wang, He, et al. (2018). "Comparative Profiling of Three ''Atheris'' Snake Venoms: ''A. squamigera, A. nitschei and A. chlorechis'' ". ''The Protein Journal'' '''37''' (4): 353–360. doi:10.1007/s10930-018-9781-y.</ref>

==Species==

{| class="wikitable sortable" |- ! Image !! Species<ref name="McD99"/>!! Taxon author<ref name="McD99"/> !!Subsp.*<ref name="ITIS"/> !! Common name !! Geographic range<ref name="McD99"/> |- | |''A. acuminata'' |Broadley, 1998 |align="center"|0 |Acuminate bush viper |Western Uganda |- | |''A. anisolepis'' |Mocquard, 1887 |align="center"|0 |Mayombe bush viper |style="width:40%"|West central Africa: Gabon, Congo, west DR Congo, north Angola |- | |''A. barbouri'' |(Loveridge, 1930) |align="center"|0 |Barbour's short-headed viper, Uzungwe Mountain bush viper |The Udzungwa and Ukinga mountains in southern Tanzania |- | |''A. broadleyi'' |D. Lawson, 1999 |align="center"|0 |Broadley's bush viper<ref>Beolens, Bo; Watkins, Michael; Grayson, Michael (2011). ''The Eponym Dictionary of Reptiles''. Baltimore: Johns Hopkins University Press. xiii + 296 pp. {{ISBN|978-1-4214-0135-5}}. (''Atheris broadleyi'', p. 39).</ref> |Cameroon, Central African Republic, Nigeria, Congo |- |120px |''A. ceratophora'' |F. Werner, 1895 |align="center"|0 |Usambara eyelash viper |The Usambara and Uzungwe Mountains in Tanzania |- |120px |''A. chlorechis''T |(Pel, 1851) |align="center"|0 |West African bush viper |West Africa including Guinea-Bissau, Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Benin, isolated locations in Nigeria, Cameroon, Equatorial Guinea, and Gabon |- | |''A. desaixi'' |Ashe, 1968 |align="center"|0 |Mount Kenya bush viper, Ashe's bush viper |Two isolated populations in Kenya: in the forests at Chuka, south-eastern Mount Kenya, and Igembe in the northern Nyambeni range |- | |''A. hetfieldi'' <ref>{{cite news |last1=Maciel |first1=Ana Rita |title=Investigadores portugueses dão o nome do vocalista dos Metallica a nova espécie de víbora africana |url=https://www.publico.pt/2020/09/05/ciencia/noticia/investigadores-portugueses-dao-nome-vocalista-metallica-nova-especie-vibora-africana-1929999 |access-date=5 September 2020 |publisher=Público |date=2020-09-05}}</ref><ref>{{Cite web|url=https://consequence.net/2020/08/venomous-snake-species-metallica-james-hetfield/|title=New venomous snake species named after Metallica's James Hetfield|date=August 28, 2020}}</ref> |Ceríaco, Marques & Bauer, 2020 |align="center"|0 |Hetfield’s bush viper | Bioko island, Equatorial Guinea |- | |''A. hirsuta'' |R. Ernst & Rödel, 2002 |align="center"|0 |Tai hairy bush viper |Ivory Coast |- |120px |''A. hispida'' |Laurent, 1955 |align="center"|0 |African hairy bush viper |Central Africa: DR Congo, south-west Uganda, west Kenya |- | |''A. katangensis'' |de Witte, 1953 |align="center"|0 |Katanga mountain bush viper |Restricted to Upemba National Park, Shaba Province in eastern DR Congo |- | |''A. mabuensis'' |Branch & Bayliss, 2009<ref>{{cite journal |vauthors=Branch WR, Bayliss J |title=A new species of ''Atheris'' (Serpentes: Viperidae) from northern Mozambique |journal=Zootaxa |volume=2113 |pages=41–54 |year=2009|doi=10.11646/zootaxa.2113.1.2 }}</ref> |align="center"|0 |Mount Mabu forest viper |Mount Mabu and Mount Namuli, northern Mozambique |- | |''A. matildae'' |Menegon, Davenport & Howell, 2011 |align="center"|0 |Matilda's horned viper |south west Tanzania |- | |''A. mongoensis'' |Collet & J.-F. Trape, 2020 |align="center"|0 |Mongo hairy bush viper |Democratic Republic of Congo |- |120px |''A. nitschei'' |Tornier, 1902 |align="center"|0 |Great Lakes bush viper |Central Africa from east DR Congo, Uganda and west Tanzania southward to north Malawi and north Zambia. |- | |''A. rungweensis'' |Bogert, 1940 |align="center"|0 |Mt. Rungwe bush viper |southwestern Tanzania, northeastern Zambia, northern Malawi |- |120px |''A. squamigera'' |(Hallowell, 1856) |align="center"|0 |Variable bush viper |West and central Africa: Ivory Coast and Ghana, eastward through southern Nigeria to Cameroon, southern Central African Republic, Gabon, Congo, DR Congo, northern Angola, Uganda, Tanzania (Rumanika Game Reserve), western Kenya, and Bioko Island |- | |''A. subocularis'' |Fischer, 1888 |align="center"|0 | |Cameroon |- |} *(''Not including the nominate subspecies'' ''T'') Type species

==Taxonomy== Other species may be encountered in literature, such as:<ref name="NRDB">{{NRDB genus|genus=Atheris|date=2 August|year=2007}}</ref><ref name="WOA">{{WOA|page=index|subject=Home|date=8 September|year=2007}}</ref>

Until relatively recently, the following species, all of which are terrestrial, were also included in the genus ''Atheris'':<ref name="Mal03"/>

*''Montatheris hindii'' (Boulenger, 1910) – montane viper *''Proatheris superciliaris'' (W. Peters, 1855) – lowland viper

Together with ''Atheris'', these three genera are sometimes referred to as the tribe Atherini.<ref name="WOA"/>

==References== {{Reflist}}

==Further reading== {{Commons}} {{Refbegin}} *{{cite journal |author=Bonaparte CL |author-link=Charles Lucien Bonaparte |title=On the Lorine genus of Parrots, ''Eclectus'', with the description of a new species, ''Eclectus cornelia'' |journal=Proceedings of the Zoological Society of London |volume=17 |pages=142–146 [145, footnote] |year=1849}} *Boulenger GA (1896). ''Catalogue of the Snakes in the British Museum (Natural History). Volume III., Containing the ...Viperidæ''. London: Trustees of the British Museum (Natural History). (Taylor and Francis, printers.) xiv + 727 pp. + Plates I–XXV. (Genus ''Atheris'', p. 508.) *{{cite journal |doi=10.1080/21564574.1996.9649964 |author=Broadley DG |title=A review of the tribe Atherini (Serpentes: Viperidae), with the descriptions of two new genera |journal=African Journal of Herpetology |volume=45 |issue=2 |pages=40–48 |year=1996|bibcode=1996AfJH...45...40B |author-link=Donald George Broadley }} *{{cite journal |author=Cope ED |author-link=Edward Drinker Cope |title=Notes upon some REPTILES of the Old World |journal=Proceedings of the Academy of Natural Sciences of Philadelphia |volume=14 |pages=337–344 [343–344] |year=1862}} *{{cite journal |author=Freed P |author-link=Paul Freed|title=''Atheris chlorechis'' (West African bush viper) |journal=Herpetological Review |volume=17 |issue=2 |pages=47–48 |year=1986}} *{{cite journal |author=Günther ACLG |author-link=Albert Günther |title=On new species of snakes in the collection of the British Museum |journal=Annals and Magazine of Natural History, Third Series |location=London |volume=11 |issue=3 |pages=20–25 [25] |year=1863|doi=10.1080/00222936308681373 }} *{{cite journal |vauthors=Lanoie L, Branch W |title=''Atheris squamiger'' : fatal envenomation |journal=Journal of the Herpetological Association of Africa |volume=39 |pages=29 |year=1991}} *{{cite journal |author=Love W |title=Bush vipers (''Atheris''): Experiences in breeding and maintenance |journal=Vivarium |volume=1 |issue=3 |pages=22–25 |year=1988}} *{{cite journal |author=Pareti KS |title=Cannibalism in a captive West African bush viper (''Atheris chloroechis'') |journal=Herpetological Review |volume=25 |issue=1 |page=17 |year=1994}} *{{cite book |author=Pitman CRS |author-link=Charles Pitman (game warden) |title=A Guide to the Snakes of Uganda |publisher=Codicote, Wheldon & Wesley |location=London |year=1974 |isbn=0-85486-020-7}} 290 pp. {{Refend}}

Category:Atheris Category:Snake genera Category:Taxa named by Edward Drinker Cope