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© Rune Midtgaard


Taxonomy of the family Viperidae
Bibliography of the genus Gloydius
Biodiversity of the family Viperidae








Genus
Gloydius

Mamushis, Asian Moccasins, (Asian Copperheads)

Mamushis, Halysottern

Mamushier

1822 Trigonocephalus Kuhl & van Hasselt [not Trigonocephalus Oppel 1811 (Serpentes)] (type species: Coluber halys Pallas 1776)
1826 Trigonocephalus Boie [not Trigonocephalus Oppel 1811 (Serpentes); not Trigonocephalus Kuhl & van Hasselt 1822 (Serpentes)] (type species: Coluber Halys Pallas 1776)
1849 Halys Gray [not Halys Fabricius 1803 (Insecta)] (type species: Coluber halys Pallas 1776)
1981 Gloydius Hoge & Romano Hoge [substitute name for Halys Gray 1849] (type species: Coluber halys Pallas 1776)
Contents: 23 species, of which 14 (60.9%) are endemic.
Remarks: Previously included in Agkistrodon (e.g., Gloyd & Conant 1990). Validity as a separate genus confirmed by several subsequent works (Knight, Densmore & Rael 1992; Parkinson, Moody & Ahlquist 1996; Cullings, Morafka, Hernandez & Roberts 1997; Vidal, Lecointre, Vie & Gasc 1997). Validity of several nominal species is still unresolved, and various authorities have suggested different arrangements.
Distribution: SW., C., and E. Asia.
Reported from: Afghanistan, Azerbaijan, Bhutan, China (Anhui, Beijing, Chongqing, Fujian, Gansu, Guangdong, Guizhou, Hebei, Heilongjiang, Henan, Hubei, Hunan, Inner Mongolia, Jiangsu, Jiangxi, Jilin, Liaoning, Ningxia, Qinghai, Shandong [incl. Daheishan Islands], Shaanxi, Shanxi, Sichuan, Tibet, Xinjiang, Yunnan, Zhejiang), India (Haryana, Himachal Pradesh, Jammu & Kashmir, Punjab, Uttar Pradesh, Uttarakhand, West Bengal), Iran, Japan (Hokkaido, Honshu, Kyushu, Shikoku, Tsushima Islands, Izu Islands, Ryukyu Islands [incl. Satsunan Islands]), Kazakhstan, Kyrgyzstan, Mongolia, Nepal, North Korea, Pakistan, Russia (incl. Kuril Islands), South Korea, Tajikistan, Turkmenistan, Uzbekistan.

Gloydius angusticeps

Narrow-headed Mamushi

Schmalkopf-Mamushi

Smalhovedet Mamushi

2018 Gloydius angusticeps Shi, Yang, Zhang, Peng, Orlov, Jiang, Ding, Hou, Huang, Huang & Li

Distribution: China (Gansu, Qinghai, Sichuan).





Gloydius blomhoffii

Japanese Mamushi, Common Mamushi

Japanische Mamushi

Japansk Mamushi

1826 Trigonocephalus Blomhoffii Boie
Halys blomhoffii Peters 1862
Ancistrodon blomhoffii Boulenger 1896
Agkistrodon blomhoffii Stejneger 1907
Ancistrodon halys blomhoffii Nikolsky 1916
Gloydius blomhoffii Hoge & Romano Hoge 1981
1849 Trigonocephalus affinis Gray (status uncertain, syn. Gloyd & Conant 1990)
Agkistrodon blomhoffii ? affinis Stejneger 1907
Agkistrodon halys affinis Mell 1929
Agkistrodon affinis Gloyd & Conant 1990
1860 Trigonocephalus blomhoffii var. megaspilus Cope (Golay & al. 1993)

Remarks: See remarks under halys and brevicaudus. Older records from Tsushima Island (Japan) refer to tsushimaensis (Gumprecht, Tillack, Orlov, Captain & Ryabov 2004). Erroneously recorded from Sakhalin Island, Russia (e.g., Gloyd & Conant 1990; David & Ineich 1999) and Taiwan (e.g., Zhao, Jiang & Huang 1979) (Zhao & Adler 1993; Gumprecht, Tillack, Orlov, Captain & Ryabov 2004).
Distribution: Japan (Hokkaido, Honshu, Kyushu, Shikoku, Izu Islands, Ryukyu Islands [incl. Satsunan Islands]), Russia (Kuril Islands).


Gloydius brevicaudus

Short-tailed Mamushi, Korean Mamushi

Kurzschwanz-Mamushi

Korthalet Mamushi

1907 Agkistrodon blomhoffii brevicaudus Stejneger
Ancistrodon blomhoffii brevicaudus Nikolsky 1914
Gloydius blomhoffi brevicaudus Hoge & Romano Hoge 1981
Gloydius brevicaudus Zhou, Zhang, Huang, Chen, Chen & Yao 2001
1977 Agkistrodon blomhoffii dubitatus Gloyd (Zhao & Adler 1993; see remarks)
Gloydius blomhoffi dubitatus Hoge & Romano-Hoge 1981
1977 Agkistrodon blomhoffii siniticus Gloyd (Wallach, Williams & Boundy 2014)
Gloydius blomhoffii siniticus Hoge & Romano Hoge 1981

Other common names:
dubitatus: Tung Ling Mamushi, (Northern Chinese Mamushi)
siniticus: Yangtze Mamushi, (Chinese Mamushi)
Remarks: Previously included cognatus and caraganus. Formerly regarded as a synonym or subspecies of blomhoffii by some authors (e.g., McDiarmid, Campbell & Touré 1999; Watkins-Colwell & Leenders 2002) or as a valid species by others (e.g., Vogel 2006; Wallach, Williams & Boundy 2014). Some authors regard dubitatus as a synonym of blomhoffii (e.g., Wallach, Williams & Boundy 2014). Herein, dubitatus is assigned to brevicaudus, following Zhao & Adler (1993), thus restricting blomhoffii to Japan. Gumprecht, Tillack, Orlov, Captain & Ryabov (2004) did not mention dubitatus in the synonymy of blomhoffii.
Distribution: China (Anhui, Beijing, Chongqing, Fujian, Guangdong [incl. Nan Ao Island], Guizhou, Hebei, Heilongjiang, Henan, Hubei, Hunan, Inner Mongolia, Jiangsu, Jiangxi, Jilin, Liaoning, Shandong, Shaanxi, Sichuan, Yunnan, Zhejiang), North Korea, South Korea.


Gloydius caraganus

Karaganda Mamushi, Karaganda Pitviper, (Siberian Pitviper)

Karaganda-Mamushi

Karaganda-mamushi

1831 Trigonocephalus caraganus Eichwald
Agkistrodon halys caraganus Chernov 1934
Ancistrodon halys caraganus Terentjev & Chernov 1949
Gloydius halys caraganus Hoge & Romano Hoge 1981
Gloydius caraganus Shi, Yang, Zhang, Qi, Li & Ding 2016
1931 Ancistrodon halys paramonovi Nikolsky (Golay & al. 1993)

Remarks: Revalidated as a separate species by Shi, Yang, Zhang, Qi, Li & Ding (2016). Validity supported by Shi, Wang, Chen, Fang, Ding, Huang, Hou, Liu & Li (2017). See remarks under halys regarding taxonomic instability of the names caraganus and halys.
Distribution: Kazakhstan, Kyrgyzstan, Mongolia, Russia, Tajikistan, Turkmenistan, Uzbekistan.


Gloydius caucasicus

Caucasian Mamushi, Caucasian Pitviper

Kaukasische Mamushi

Kaukasisk Mamushi

1916 Ancistrodon halys caucasicus Nikolsky
Agkistrodon halys caucasicus Chernov 1934
Agkistrodon intermedius caucasicus Gloyd & Conant 1982
Gloydius halys caucasicus Hoge & Romano Hoge 1981
Gloydius caucasicus Asadi, Montgelard, Nazarizadeh, Moghaddasi, Fatemizadeh, Simonov, Kami & Kaboli 2019

Remarks: Previously regarded as a subspecies of halys (e.g., David & Vogel 2010; Midtgaard 2019).
Distribution: Afghanistan, Azerbaijan, Iran, Turkmenistan.





Gloydius changdaoensis

Jiaodong Mamushi

Jiaodong-Mamushi

Jiaodong-mamushi

1999 Gloydius saxatilis changdaoensis Li
Gloydius changdaoensis Shi, Wang, Chen, Fang, Ding, Huang, Hou, Liu & Li 2017

Distribution: China (Jiangsu, Shandong [incl. Daheishan Islands]).


Gloydius cognatus

Alashan Mamushi, Alashan Pitviper

Alashan-Mamushi

Alashan-mamushi

1977 Agkistrodon halys cognatus Gloyd (David & Vogel in Visser 2015)
Gloydius halys cognatus Hoge & Romano Hoge 1981
Gloydius cognatus Shi, Yang, Zhang, Qi, Li & Ding 2016

Remarks: Previously regarded as a synonym of brevicaudus (e.g., David & Vogel in Visser 2015) or a subspecies of halys (e.g., Hoge & Romano Hoge 1981). Revalidated as a separate species by Shi, Yang, Zhang, Qi, Li & Ding (2016). Validity supported by Shi, Wang, Chen, Fang, Ding, Huang, Hou, Liu & Li (2017).
Distribution: China (Gansu, Inner Mongolia, Ningxia, Qinghai, Sichuan).


Gloydius halys

Central Asian Mamushi, Asiatic Pitviper, Asiatic Moccasin, (Halys' Pitviper, Pallas' Pitviper)

Halysschlange, Schitomordnik

Centralasiatisk Mamushi, Halysslange

1776 Coluber Halys Pallas
Trigonocephalus halys Lichtenstein in Eversman & Lichtenstein 1823
Vipera halys Lichtenstein in Eversman & Lichtenstein 1823 (as cited in the synonymy of T. halys)
Ancistrodon halys Boulenger 1896
Agkistrodon halys Stejneger 1907
Gloydius halys Hoge & Romano Hoge 1981
1820 Vipera (Echidna) aspis pallasii Merrem (McDiarmid, Campbell & Touré 1999)
1876 Halys pallasii Blanford [not Vipera (Echidna) aspis pallasii Merrem 1820] (Golay & al. 1993)
1933 Ancistrodon halys persicus Rendahl (Wallach, Williams & Boundy 2014)
Agkistrodon halys persicus Golay & al. 1993
1983 Agkistrodon halys boehmei Nilson
1993 Agkistrodon halys mogoi Bour (Ananjeva, Munkhbayar, Orlov, Orlova, Semenov & Terbisk 1997; Orlov & Barabanov 1999)
Gloydius halys mogoi David & Vogel 2010
2017 Gloydius halys ubsunurensis Kropachev & Orlov

Other common names:
boehmei: Afghan Mamushi, Afghan Pitviper, (Böhme’s Pitviper)
halys (incl. mogoi): Mongolian Mamushi, (Siberian Pitviper)
ubsunurensis: Ubsunur Mamushi, Ubsunur Pitviper
Remarks: Previously included blomhoffi, caraganus, caucasicus, intermedius, and stejnegeri. Confusion or lack of consensus still prevails regarding the use of the names halys and caraganus. The latter was considered a distinct species by Shi, Yang, Zhang, Qi, Li & Ding (2016), as already suspected to be by Gumprecht, Tillack, Orlov, Captain & Ryabov (2004), which is followed herein. However, Bour (1993), followed by some other authors (e.g., David 2007; David & Vogel 2010), regarded halys as the correct name for the western species referred to herein as caraganus (above), and he introduced the new name Agkistrodon halys mogoi for the eastern form, referred to herein as Gloydius h. halys. This taxonomic instability is related to uncertainty regarding the type locality. Orlov & Barabanov (2000) selected a neotype from Siberia, an action critisized by David (2007) and David & Vogel (2010) with reference to Bour (1993), who presented evidence that the type was collected north of Astrakhan, near the border between Russia and Kazakhstan. Pending the establishment of a consensus regard these taxonomic issues, the work of Orlov & Barabanoc (1999) is provisionally followed herein.
Distribution: Afghanistan, Azerbaijan, China (Inner Mongolia), Iran, Kazakhstan, Kyrgyzstan, Mongolia, Russia, Tajikistan, Turkmenistan, Uzbekistan.


Gloydius himalayanus

Himalayan Mamushi, Brown Himalayan Viper, (Himalayan Pitviper)

Himalaya-Mamushi

Himalaya-mamushi, (Himalayagrubeorm)

1864 Halys himalayanus Günther
Trigonocephalus himalayanus Strauch 1873
Crotalus halus himalayanus Higgins 1873
Ancistrodon himalayanus Boulenger 1890
Agkistrodon himalayanus Hatta 1928
Gloydius himalayanus Hoge & Romano Hoge 1981

Remarks: Presence in Sikkim, India, needs confirmation (Gumprecht, Tillack, Orlov, Captain & Ryabov 2004), although likely, since the species is found both to the east and to the west of the state. A single specimen has been reported from Afghanistan, but was later reidentified as Macrovipera lebetina (Wagner, Bauer, Leviton, Wilms & Böhme 2016). Records from Bangladesh (e.g., Sarker & Sarker 1985) are erroneous (Gumprecht, Tillack, Orlov, Captain & Ryabov 2004).
Distribution: Bhutan, India (Haryana, Himachal Pradesh, Jammu & Kashmir, Punjab, Uttar Pradesh, Uttarakhand, West Bengal), Nepal, Pakistan.


Gloydius huangi

Lancang Mamushi, Lancang Plateau Viper

Lancang-Mamushi

Lancang-mamushi

2019 Gloydius huangi Wang, Ren, Dong, Jiang, Shi, Siler & Che

Distribution: China (Tibet).





Gloydius intermedius

Siberian Mamushi, (Central Asian Pitviper, Amur Pitviper)

Zentralasiatische Mamushi

Siberisk Mamushi

1868 Trigonocephalus intermedius Strauch
Halys intermedius Peters 1877
Ancistrodon intermedius Boulenger 1896
Agkistrodon blomhoffii intermedius Stejneger 1907
Ancistrodon blomhoffii intermedius Despax 1913
Ancistrodon halys intermedius Nikolsky 1916
Agkistrodon halys intermedius Stejneger 1925
Agkistrodon intermedius Werner 1929
Gloydius halys intermedius Hoge & Romano Hoge 1981
Gloydius intermedius Kraus, Mink & Brown 1996

Remarks: Previously included stejnegeri. Formerly regarded as a subspecies of blomhoffii (e.g., Stejneger 1907), or of halys (e.g., Nikolsky 1916). Some authors have considered saxatilis a synonym of intermedius.
Distribution: China (Gansu, Heilongjiang, Inner Mongolia, Ningxia, Shaanxi, Shanxi, Xinjiang), Russia.


Gloydius lijianlii

Shandong Mamushi

Shandong-Mamushi

Shandong-mamushi

2009 Gloydius lijianlii Jiang & Zhao

Distribution: China (Shandong: Daheishan Islands).


Gloydius liupanensis

Liupan Mamushi

Liupan-Mamushi

Liupan-mamushi

1989 Agkistrodon halys liupanensis Liu, Song & Luo
Gloydius liupanensis Xu, Liu, Myers, Wang, Huang, He, Peng & Guo 2012

Distribution: China (Gansu, Ningxia).





Gloydius monticola

Likiang Mamushi, Likiang Pitviper, Mountain Mamushi, Yunnan Mamushi

Likiang-Mamushi

Likiang-mamushi

1922 Ancistrodon blomhoffii monticola Werner
Agkistrodon halys monticola Gee 1929
Agkistrodon monticola Mell 1929
Ancistrodon halys monticola Rendahl 1933
Agkistrodon blomhoffi monticola Mertens 1934
Gloydius monticola Hoge & Romano Hoge 1981
Gloydius strauchi monticola David & Ineich 1999
1931 Agkistrodon strauchi popei Mell "1929" (Golay & al. 1993; Zhao & Adler 1993)

Remarks: Considered a synonym of strauchi by Zhao (1995), but as a valid species in most recent publications (e.g., McDiarmid, Campbell & Touré 1999, Orlov, Ananjeva, Barabanov, Ryabov & Kha-Likov 2002, Gumprecht, Tillack, Orlov, Captain & Ryabov 2004, Vogel 2006).
Distribution: China (Yunnan).


Gloydius qinlingensis

Shaanxi Mamushi

Shaanxi-Mamushi

Shaanxi-mamushi

1985 Agkistrodon halys qinlingensis Song & Chen
Gloydius qinlingensis Guo, Zhang & Chen 1999

Distribution: China (Shaanxi).





Gloydius rickmersi

Kyrgyz Mamushi

Kirgisische Mamushi

Kirgisisk Mamushi

2015 Gloydius rickmersi Wagner, Tiutenko, Borkin & Simonov

Remarks: Validity supported by Shi, Wang, Chen, Fang, Ding, Huang, Hou, Liu & Li (2017).
Distribution: Kyrgyzstan.


Gloydius rubromaculatus

Red-spotted Mamushi, Red-spotted Alpine Pitviper, Tongtianhe Pitviper

Rotfleckige Mamushi

Rødplettet Mamushi

2017 Gloydius rubromaculatus Shi, Wang, Chen, Fang, Ding, Huang, Hou, Liu & Li

Distribution: China (Qinghai, Sichuan, Tibet).


Gloydius saxatilis

Rock Mamushi, Brown Mamushi

Felsen-Mamushi

Klippe-mamushi

1937 Ancistrodon saxatilis Emelianov
Agkistrodon saxatilis Gloyd 1972
Agkistrodon intermedius saxatilis Gloyd & Conant 1982
Gloydius saxatilis Hoge & Romano Hoge 1981
1980 Agkistrodon shedaoensis continentalis Zhao (Golay & al. 1993)

Remarks: Previously included changdaoensis. Formerly regarded as a synonym or subspecies of intermedius (e.g., Gloyd & Conant 1982, 1990; Orlov & Barabanov 2000; Gumprecht, Tillack, Orlov, Captain & Ryabov 2004; David & Vogel in Visser 2015). Treated as a valid species in most recent publications (e.g., Zhao & Adler 1993; Golay & al. 1993; McDiarmid, Campbell & Touré 1999; Xu, Liu, Myers, Wang, Huang, He, Peng & Guo 2012; Wallach, Williams & Boundy 2014).
Distribution: China (Heilongjiang, Inner Mongolia, Jiangsu, Jilin, Liaoning, Shandong [incl. Daheishan Islands]). North Korea, Russia, South Korea.


Gloydius shedaoensis

Shedao Island Mamushi, Shedao Island Pitviper

Shedao-Mamushi

Shedao-mamushi

1979 Agkistrodon shedaoensis Zhao
Agkistrodon intermedius shedaoensis Welch 1994
Gloydius shedaoensis McDiarmid, Campbell & Touré 1999
Gloydius saxatilis shedaoensis Zhou, Zhang, Huang, Chen, Chen & Yao 2001
1999 Gloydius shedaoensis qianshanensis Li

Distribution: China (Liaoning [Shedao Island and mainland]).


Gloydius stejnegeri

Gobi Mamushi, Gobi Pitviper, (Northern Chinese Pitviper)

Gobi-Mamushi

Gobi-mamushi

1933 Ancistrodon halys stejnegeri Rendahl
Agkistrodon intermedius stejnegeri Gloyd & Conant 1982
Gloydius halys stejnegeri Gumprecht, Tillack, Orlov, Captain & Ryabov 2004
Gloydius stejnegeri Shi, Yang, Zhang, Qi, Li & Ding 2016

Remarks: Previously regarded as a synonym of intermedius (e.g., David & Vogel 2010; David & Vogel in Visser 2015). Revalidated as a separate species by Shi, Yang, Zhang, Qi, Li & Ding (2016). Validity supported by Shi, Wang, Chen, Fang, Ding, Huang, Hou, Liu & Li (2017).
Distribution: China (Beijing, Hebei, Inner Mongolia, Shaanxi, Shanxi).


Gloydius strauchi

Sichuan Mamushi (Tibetan Mamushi, Strauch’s Pitviper, Strauch's Mamushi)

Sichuan Mamushi, (Strauchs Grubenotter)

Sichuan-mamushi, (Tibetansk Mamushi)

1912 Ancistrodon Strauchi Bedriaga
Agkistrodon strauchi Stejneger 1925
Ancistrodon halys strauchi Pavlov 1932
Gloydius strauchi Hoge & Romano Hoge 1981
1912 Agkistrodon tibetanus Barbour (Golay & al. 1993; Zhao & Adler 1993)
Ancistrodon tibetanus Werner 1922

Remarks: Previously included qinlingensis and liupanensis, as well as the populations now assigned to angusticeps and rubromaculatus (Shi, Yang, Zhang, Peng, Orlov, Jiang, Ding, Hou, Huang, Huang & Li 2018). Exact western limit of distribution is unclear, since some records may refer to rubromaculatus or angusticeps (Shi, Wang, Chen, Fang, Ding, Huang, Hou, Liu & Li 2017; Shi, Yang, Zhang, Peng, Orlov, Jiang, Ding, Hou, Huang, Huang & Li 2018). A single report from Yunnan (He & Zhou 2002) with unknown locality.
Distribution: China (Sichuan, Yunnan).


Gloydius tsushimaensis

Tsushima Mamushi, Tsushima Island Pitviper

Tsushima-Mamushi

Tsushima-mamushi

1994 Agkistrodon tsushimaensis Isogawa, Moriya & Mitsui
Gloydius tsushimaensis Gumprecht, Tillack, Orlov, Captain & Ryabov 2004

Distribution: Japan (Tsushima Islands).


Gloydius ussuriensis

Ussuri Mamushi, Ussurian Pitviper, (Amur Mamushi)

Ussuri-Mamushi

Ussuri-mamushi

1929 Ancistrodon blomhoffii ussuriensis Emelianov
Agkistrodon blomhoffii ussuriensis Chernov 1934
Agkistrodon halys ussuriensis Klemmer 1963
Gloydius halys ussuriensis Hoge & Romano Hoge 1981
Agkistrodon ussuriensis Toriba 1986
Gloydius ussuriensis Vogel 2006
1972 Agkistrodon caliginosus Gloyd (Toriba 1988; McDiarmid, Campbell & Touré 1999; Gumprecht, Tillack, Orlov, Captain & Ryabov 2004)
Gloydius caliginosus Hoge & Romano Hoge 1981

Other common names:
caliginosus: Dusky Mamushi, Caliginous Mamushi, Kankoku Mamushi
Distribution: China (Heilongjiang, Inner Mongolia, Jilin, Liaoning), Mongolia, North Korea, Russia, South Korea.