After Gonrad, Ast, Montanari & Norell, 2011 (fig. 27), and Conrad, Balcarcel & Mehling, 2012, with untreated Recent species from Uetz, Etzold & Chenna, 1995-2000 [“The Reptile Database” (former EMBL Reptile Database)] and Wikipedia
<==o Varanus Merrem, 1820 [Hydrosaurus Wagler, 1830; Megalania; Notiosaurus; Odantria] L.Eoc. As. Af., Olig.-R. Aust., M.Mioc.-R. As.; L.Mioc.-R. Af., Mioc. Eu. (varaanit; monitor lizards) |?- V. sp. [Holmes et al., 2010] [Late Eocene-Early Oligocene, Northeastern Africa (Fayum)] |--o (Polydaedalus) | |-- †V. rusingensis Clos, 1995 | |-- V. ornatus (Daudin, 1803) (Ornate Monitor) | `--+--o V. albigularis (Daudin, 1802) sensu Conrad et al., 2008 (White-throated Monitor, Rock Monitor; valkokurkkuvaraani) | | |-- V. a. albigularis (White-throated Monitor) | | |-- V. a. angolensis (Angola White-throated Monitor) | | `-- V. a. microstictus (Eastern White-throated Monitor) | |-- V. exanthematicus (Bosc, 1792) (Steppe monitor, Savannah Monitor; arovaraani) | |-- V. niloticus (Linnaeus, 1766) (Nile monitor; Qamu; niilinvaraani) | `?- V. yemenensis Böhme, Joger & Schätti, 1989 (Yemen Monitor; jemeninvaraani, arabianvaraani) `--+--o (Psammosaurus) Fitzinger, 1826 | |?- †V. darevskii Levshakova, 1986 | `--o V. griseus (Daudin, 1803) (Desert Monitor; aavikkovaraani) | |-- V. g. griseus (Desert Monitor) | |-- V. g. caspius (Caspian Monitor) | `-- V. g. koniecznyi (Thar Desert monitor) `--+-- †V. mokrensis Ivanov, Ruta, Klembara & Böhme, 2018 |-- †V. marathonensis Weithofer, 1888 [incl. Varanus atticus Nopcsa, 1908; Varanus (Varaneades) amnhophilis Conrad, Balcarcel, & Mehling, 2012 [AMNH FR 30630] sensu Villa et al., 2018] `--+-- V. kordensis (Meyer, 1874) (Biak Tree Monitor) |--+--+-- †V. hooijeri | | `--o (Philippinosaurus) | | |?- V. bitatawa Welton, Siler, Bennett, Diesmos, Ruya, Dugay, Rico, Van Weerd & Brown, 2010 (Northern Sierra Madre Forest Monitor Lizard, Bitatawa) | | |?- V. mabitang Gaulke & Curio, 2001 (Panay Monitor) | | `-- V. olivaceus Hallowell, 1857 (Gray's Monitor) | `--o (Euprepiosaurus) | |?- V. boehmei Jacobs, 2003 (Golden-spotted Tree Monitor) | |?- V. bogerti Mertens, 1950 (Bogert's Tree Monitor) | |?- V. caerulivirens Ziegler, Böhme & Philipp, 1999 (Turquoise Monitor) | |?- V. cerambonensis Philipp, Böhme & Ziegler, 1999 (Ambon Monitor) | |?- V. finschi Böhme, Horn & Ziegler, 1994 (Finsch's Monitor) | |?- V. juxtindicus Böhme, Philipp & Ziegler, 2002 (Rennell Island Monitor) | |?- V. macraei Böhme & Jacobs, 2001 (Blue-spotted Tree Monitor) | |?- V. lirungensis Koch, Arida, Schmitz, Böhme & Ziegler, 2009 (Lirung Monitor) | |?- V. obor Weijola & Sweet, 2010 (Sago Monitor) | |?- V. rainerguentheri Ziegler, Böhme & Schmitz, 2007 (Rainer Guenther's Monitor) | |?- V. reisingeri Eidenmüller & Wicker, 2005 (Reisinger's Tree Monitor) | |?- V. telenesetes Sprackland, 1991 (Rossel Island Tree Monitor) | |?- V. zugorum Böhme & Ziegler, 2005 (Zug's Monitor) | |--+-- V. prasinus (Schlegel, 1839) (Emerald Tree Monitor, Emerald Monitor) | | `--+-- V. beccari (Doria, 1874) (Black Tree Monitor) | | `-- V. keithhornei (Wells & Wellington, 1985) (Canopy Goanna, Nesbit River Monitor) | `--+--+-- V. melinus Böhme & Ziegler, 1997 (Quince Monitor) | | `-- V. indicus (Daudin, 1802) (Mangrove Monitor, Indian monitor; intianvaraani) | `--+-- V. jobiensis Ahl, 1932 (Peach-throat Monitor) | `--+-- V. doreanus (Meyer, 1874) (Blue-tailed Monitor) | `-- V. yuwonoi Harvey & Barker, 1998 (Tri-colored Monitor) `--+--+--+-- V. (Empagusia) flavescens (Hardwicke & Gray, 1827) | | `--+-- †V. (Varaneades) amnhophilis Conrad, Balcarcel, & Mehling, 2012 [AMNH FR 30630] | | `--+-- V. (Tectovaranus) dumerilii (Schlegel, 1839) | | `--o (Indovaranus) | | |-- V. nebulosus (Gray, 1931) [V. bengalensis nebulosus] (Clouded Monitor) | | `-- V. bengalensis (Daudin, 1802) (Bengal monitor; common Indian monitor; bengalinvaraani) | `--+-- V. (Dendrovaranus) rudicollis (Gray, 1845) | `--o (Soterosaurus) | |?- V. nuchalis (Günther, 1872) (Large-scaled Water Monitor) | |?- V. palawanensis Koch, Gaulke & Böhme, 2010 (Palawan Water Monitor) | |?- V. rasmusseni Koch, Gaulke & Böhme, 2010 (Rasmussen's Water Monitor) | |-- V. cumingi Martin, 1839 (Cuming's Water Monitor) | `--+-- V. marmoratus (Wiegmann, 1834) (Phiippine Water Monitor) | `--+-- V. togianus (Peters, 1872) (Togian Water Monitor) | `--o V. salvator (Laurenti, 1768) Laurenti, 1768 (Water Monitor; juovavaraani) | |-- V. s. salvator (Sri Lankan Water Monitor) | |-- V. s. andamanensis (Andaman Islands Water Monitor) | |-- V. s. bivittatus (Two-striped Water Monitor) | |-- V. s. macromaculatus (Southeast Asian Water Monitor) | `-- V. s. ziegleri (Ziegler's Water Monitor) `--+--o (Varanus) Merrem, 1820 | |--+-- V. spenceri Lucas & Frost, 1903 (Spencer's Goanna) | | `--+-- V. (Papusaurus) salvadorii (Peters & Doria, 1878) (krokotiilivaraani) | | `--+-- V. komodoensis Ouwens, 1912 (Komodo dragon; komodonvaraani) | | |-- †V. priscus (Owen, 1859) [Megalania prisca Owen, 1859; Notiosaurus] sensu Conrad et al., 2008 (megalania) | | `-- V. varius (White, 1790) (Lace Monitor; kirjovaraani) | `--+--+-- V. mertensi Glauert, 1951 (Mertens' Monitor; mertensinvaraani) | | `-- V. giganteus (Gray, 1845) (Perentie) | `--+-- V. rosenbergi Mertens, 1957 (Rosenberg's Goanna, Heath Monitor) | `--+--o V. panoptes Storr, 1980 (suurvaraani) | | |-- V. p. panoptes (Argus Monitor) | | |-- V. p. horni (Horn's Monitor) | | `-- V. p. rubidus (Yellow-spotted Monitor) | `--o V. gouldii (Gray, 1838) (Sand Goanna, Gould's Goanna; gouldinvaraani) | |-- V. g. gouldii | `-- V. g. flavirufus [V. flavirufus] `--o (Odatria) (kääpiövaraanit; dwarf monitors) |?- V. auffenbergi Sprackland, 1999 (Peacock Monitor) |?- V. bushi Aplin, Fitch & King, 2006 (Pilbara Monitor, Bush's Monitor) |?- V. caudolineatus Boulenger, 1885 (Stripe-tailed Monitor) |?- V. similis Mertens, 1958 (Spotted Tree Monitor) |?- V. glauerti Mertens, 1957 (Kimberley Rock Monitor) |--+-- V. semiremex Peters, 1869 (Rusty Monitor) | |--+-- V. glebopalma Mitchell, 1955 (Black-palmed Rock Monitor) | | `-- V. pilbarensis Storr, 1980 (Pilbara Rock Monitor) | `--+--o V. tristis (Schlegel, 1839) (Black-headed Monitor) | | |-- V. t. tristis | | `-- V. t. orientalis (Freckled Monitor) | `--+-- V. scalaris Mertens, 1941 (Banded Tree Monitor) | `--+-- V. timorensis Gray, 1831 (Timor dwarf monitor; timorinkääpiövaraani) | `-- V. mitchelli Mertens, 1958 (Mitchell's Water Monitor) `--+--+-- V. brevicauda Boulenger, 1898 (Short-tailed Monitor; lyhythäntävaraani) | `-- V. eremius Lucas & Frost, 1895 (Desert Pygmy Monitor) `--+-- V. gilleni Lucas & Frost, 1895 (Pygmy Mulga Monitor) `--+--+-- V. kingorum Storr, 1980 (King's Monitor) | `-- V. primodius Mertens, 1942 (Blunt-spined Monitor) `--+--o V. storri Mertens, 1966 (Storr's Monitor) | |-- V. s. storri (Eastern Storr's Monitor) | `-- V. s. ocreatus (Western Storr's Monitor) `--+-- V. baritji King & Horner, 1987 (White's Dwarf Monitor, Black-spotted Ridge-tailed Monitor, Black-spotted Spiny-tailed Monitor) `--o V. acanthurus Boulenger, 1885 (Ridge-tailed Monitor) |-- V. a. acanthurus (Spiny-tailed Monitor) |-- V. a. brachyurus (Common Ridge-tailed Monitor) `-- V. a. insulanicus (Island Ridge-tailed Monitor)
Untreated Varanus-species:
†V. catalaunicus (nomen dubium)
†V. deserticolus Bolkay, 1913 (nomen dubium)
†V. hofmanni Roger, 1898 (nomen dubium)
†V. lungui Zerova & Chkhikvadze, 1986 (nomen dubium)
†V. semjonovi Zerova & Chkhikvadze, 1986 (nomen dubium)
†V. pronini Zerova & Ckhikvadze, 1986 (nomen dubium)
†V. tyrasiensis Zerova & Ckhikvadze, 1986 (non? Lungu et al., 1983) (nomen dubium)
†V. lungui Zerova & Chkhikvadze, 1986 (nomen dubium)
†V. sivalensis Falconer, 1868
?V. karlschmidti
V. spinulosus Mertens, 1941 (Spiny-Necked Monitor, Solomon Island Spiny Monitor)
?V. teriae
Altenative phylogenies of Varanus:
After Villa et al., 2018
<==o Varanus
|--+-- †V. rusingensis
| `--+-- †V. cf. bengalensis
| `--o (Empagusia)
| |-- V. ornatus
| `--+-- V. niloticus
| `--+-- V. exanthematicus
| `-- V. albigularis
`--+-- V. griseus
`--+-- †V. mokrensis
`--+--o (Euprepiosaurus)
| |--+-- †V. hooijeri
| | `-- V. olivaceus
| `--+--+-- V. prasinus
| | `--+-- V. kordensis
| | `--+-- V. keithhornei
| | `-- V. beccarii [sic.]
| `--+--+-- V. melinus
| | `-- V. indicus
| `--+-- V. jobiensis
| `--+-- V. doreanus
| `-- V. yuwonoi
`--+--+--+-- V. rudicollis
| | `--+-- V. cumingi
| | `--+-- V. marmoratus
| | `--+-- V. togianus
| | `--+-- V. bivittatus
| | `-- V. salvator
| `--+-- V. flavescens
| `--+-- †V. marathonensis
| `--o (Indovaranus)
| |-- V. dumerilii
| `--+-- V. nebulosus
| `-- V. bengalensis
`--+--o (Varanus)
| |--+-- V. spenceri
| | `--+-- V. salvadorii
| | `--+-- V. komodoensis
| | `--+-- †V. priscus
| | `-- V. varius
| `--+--+-- V. giganteus
| | `-- V. mertensi
| `--+-- V. rosenbergi
| `--+-- V. gouldii
| `-- V. panoptes
`--o (Odatria)
|--+--+-- V. brevicauda
| | `-- V. eremius
| `--+-- V. gilleni
| `--+--+-- V. kingorum
| | `-- V. primodius
| `--+-- V. storri
| `--+-- V. acanthurus
| `-- V. baritji
`--+--+-- V. pilbarensis
| `-- V. glebopalma
`--+-- V. tristis
`--+--+-- V. scalaris
| `-- V. semiremex
`--+-- V. mitchelli
`-- V. timorensis
Reference(s):
|
After Ivanov, Ruta, Klembara & Böhme, 2018
<==o Varanus
|-- †V. amnhophilis
`--+--+-- V. varius
| `--+--+-- V. mertensi
| | `-- V. komodoensis
| `--+-- V. salvadorii
| `--+-- †V. cf. marathonensis
| `-- †V. priscus
`--+--+--+-- V. storri
| | `-- V. glebopalma
| `--+-- V. tristis
| `--+-- V. melinus
| |--+-- V. prasinus
| | `-- V. kordensis
| `--+-- V. semiremex
| `-- V. brevicauda
`--+--+-- V. timorensis
| `--+-- V. primodius
| `--+-- V. mitchelli
| `-- V. gilleni
`--+--+-- V. pilbarensis
| `-- V. eremius
`--+--+-- V. gouldii
| `--+-- V. baritji
| `-- V. acanthurus
`--+--+-- V. yuwonoi
| `--+-- V. cumingi
| `--+-- V. marmoratus
| `--+-- V. salvator
| `--+-- V. mytilini
| `--+-- †V. marathonensis
| `-- V. bivittatus
`--+-- V. giganteus
`--+--+-- V. bengalensis
| `--+-- V. griseus
| `--+-- V. nebulosus
| `-- V. dumerilii
`--+-- V. rudicollis
`--+-- †V. mokrensis
`--+-- V. flavescens
`--+--+-- V. togianus
| `--+--+-- V. indicus
| | `-- V. doreanus
| `--+-- V. kingorum
| `--+-- V. jobiensis
| `-- V. beccari
`--+-- V. exanthematicus
`--+-- V. niloticus
`--+--+-- V. albigularis
| `--+-- V. olivaceus
| `--+-- V. keithhornei
| `-- †V. cf. bengalensis
`--+--+-- †V. hooijeri
| `--+-- †V. rusingensis
| `-- V. ornatus
`--+-- V. panoptes panoptes
`--+-- V. rosenbergi
`--+-- V. spenceri
`-- V. panoptes horni
Reference(s):
|
After Ast, 2001
<==o Varanus
|--+-- V. griseus
| `--+-- V. exanthematicus
| `-- V. niloticus
`--+--+--+--+-- V. rudicollis
| | | `-- V. salvator
| | |-- V. flavescens
| | `--+-- V. dumerilii
| | `-- V. bengalensis
| `--+-- V. olivaceus
| `--+--+-- V. prasinus
| | `--+-- V. beccari
| | `-- V. keithhornei
| `--+--+-- V. melinus
| | `-- V. indicus
| `--+-- V. jobiensis
| `--+-- V. doreanus
| `-- V. yuwonoi
`--+--+-- V. salvadorii
| `--+-- V. komodoensis
| `-- V. varius
`--+--+--+-- V. spenceri
| | `-- V. mertensi
| `--+-- V. giganteus
| `--+-- V. panoptes
| `-- V. gouldii
`--o (Odatria)
|--+-- V. eremius
| `--+-- V. gilleni
| `--+--+-- V. kingorum
| | `-- V. primodius
| `--+-- V. storri
| `--+-- V. acanthurus
| `-- V. baritji
`--+--+-- V. pilbarensis
| `-- V. glebopalma
`--+--+-- V. glauerti
| `-- V. tristis
`--+-- V. scalaris
`--+-- V. semiremex
`--+-- V. mitchelli
`-- V. timorensis
Reference(s):
|
After Conrad, Rieppel & Grande, 2008 (fig. 2 “Adams consensus”)
<==o Varanus
|-- V. acanthurus
|-- V. semiremex
|--+-- V. gilleni
| `-- V. eremius
|--+-- V. kordensis
| |-- V. prasinus
| `--o V. tristis
| |-- V. t. tristis
| `-- V. t. orientalis
`--+--+-- V. rudicollis
| `--+-- V. olivaceus
| `-- V. dumerilii
`--+-- V. indicus
`--+-- V. flavescens
`--+--+-- †V. rusingensis
| `--+-- V. albigularis
| |-- V. exanthematicus
| |-- V. griseus
| `-- V. niloticus
`--+-- V. bengalensis
|-- V. gouldii
`--+-- V. salvator
`--+-- V. varius
`--+-- V. komodoensis
`--+-- V. salvadorii
`-- †V. priscus
Reference(s):
|
After Pepin, 199?, and Pianka, 199?
<==o Varanus
|--o (Odatria)
| |?- V. (O.) eremius
| |--+--+-- V. (O.) scalaris
| | | `?-+-- V. (O.) timorensis
| | | `--+-- V. (O.) mitchelli
| | | `-- V. (O.) semiremex
| | `--+?- V. (O.) glauerti
| | `--o V. (O.) tristis
| | |-- V. (O.) t. tristis
| | `-- V. (O.) t. orientalis
| `--+?- V. (O.) kingorum
| |--+?- V. (O.) primodius
| | `-- V. (O.) acanthurus
| `--+?- V. (O.) brevicauda
| `-- V. (O.) gilleni
|--+?- †V. rusingensis Clos, 1995
| |?- V. yemenensis
| `--+?- V. exanthematicus
| |-- V. griseus
| `--+-- V. albigularis
| `-- V. niloticus
`--+--+?- V. olivaceus
| |--+--+?- V. rudicollis
| | | `-- V. salvator
| | `--+?-+?- V. prasinus
| | | |?- V. beccari
| | | |?- V. bogerti
| | | `?- V. telenesetes
| | `--+?- V. indicus
| | `--+?- V. jobiensis
| | `-- V. karlschmidti
| `--+-- V. dumerilii
| `--+?- V. bengalensis
| `-- V. flavescens
`--o (Varanus)
|?- V. teriae
|-- V. (V.) salvadorii
`--+-- V. (V.) mertensi
|?- V. (V.) spenceri
`--+-- V. (V.) rosenbergi
|?- †V. (V.) prisca
|--o V. (V.) gouldii
| |-- V. (V.) g. gouldii
| `-- V. (V.) g. flavirufus
|-- V. (V.) panoptes
|-- V. (V.) giganteus
|-- V. (V.) varius
`-- V. (V.) komodoensis
|
Reference(s):
- Ast, J. C., 2001: Mitochondrial DNA Evidence and Evolution in Varanoidea (Squamata).
–Cladistics: Vol. 17, #3, pp. 211-226 [doi: 10.1111/j.1096-0031.2001.tb00118.x] - Clos, L. M., 1995: A new species of Varanus (Reptilia: Sauria) from the Miocene of Kenya.
–Journal of Vertebrate Paleontology: Vol. 15, #2, pp. 254-267 - Conrad, J. L., Ast, J. C., Montanari, S. & Norell, M. A., 2011: A combined evidence phylogenetic analysis of Anguimorpha (Reptilia: Squamata).
–Cladistics: Vol. 27, #3, pp. 230-277 [doi: 10.1111/j.1096-0031.2010.00330.x] - Conrad, J. L., Balcarcel, A. M. & Mehling, C. M., 2012: Earliest Example of a Giant Monitor Lizard (
Varanus , Varanidae, Squamata).
–Plos One: Vol. 7, #8, pp. e41767 [doi: 10.1371/journal.pone.0041767] - Conrad, J. L., Rieppel, O. C. & Grande, L., 2008: Re-assessment of varanid evolution based on new data from Saniwa ensidens Leidy, 1870 (Squamata, Reptilia).
–American Museum Novitates: #3630, pp. 1-15 - Hocknull, S. A., Piper, P. J., van den Bergh, G. D., Due, R. A., Morwood, M. J. & Kurniawan, I., 2009: Dragon´s Paradise Lost: palaeobiogeography, evolution and extinction of the largest-ever terrestrial lizards (Varanidae).
–Plos One: Vol. 4, #9, pp. e7241(1-15) [doi: 10.1371/journal.pone.0007241] - Holmes, R. B., Murray, A. M., Attia, Y. S., Simons, E. L. & Chatrath, P., 2010: Oldest known Varanus (Squamata: Varanidae) from the Upper Eocene and Lower Oligocene of Egypt: support for an African origin of the genus.
–Palaeontology: Vol. 53, #5, pp. 1099-1110 [doi: 10.1111/j.1475-4983.2010.00994.x] - Ivanov, M., Ruta, M., Klembara, J. & Böhme, M., 2018: A new species of Varanus (Anguimorpha: Varanidae) from the early Miocene of the Czech Republic, and its relationships and palaeoecology.
–Journal of Systematic Palaeontology: Vol. 16, #9, pp. 767-797 [doi: 10.1080/14772019.2017.1355338] - Pepin, D. J., 199?: The origin of monitor lizards based on a review of the fossil evidence.
–iNet: No longer available! - Pianka, E. R., 199?: Evolution of varanid lizards.
–iNet: http://uts.cc.utexas.edu/~varanus/varanus.html - Rage, J.-C. & Bailon, S., 2005: Amphibians and squamate reptiles from the late Early Miocene (MN 4) of Béon 1 (Montréal-du-Gers, southwestern France).
–Geodiversitas: Vol. 27, #3, pp. 413-441 - Uetz, P., Etzold, T. & Chenna, R. (eds.), 1995-2000: The Reptile Database [former “EMBL Reptile Database”].
–iNet: http://www.reptile-database.org/ - Villa, A., Abella, J., Alba, D. M., Almécija, S., Bolet, A., Koufos, G. D., Knoll, F., Luján, À. H., Morales, J., Robles, J. M., Sánchez, I. M. & Delfino, M., 2018: Revision of Varanus marathonensis (Squamata, Varanidae) based on historical and new material: morphology, systematics, and paleobiogeography of the European monitor lizards.
–Plos One: Vol. 13, #12, Art.nr. e0207719, pp. 1-46 [doi: 10.1371/journal.pone.0207719]