After Ast, 2001, 2002b (from Conrad et al., 2008, fig.5), with untreated Recent species from Uetz, Etzold & Chenna, 1995-2000 [“The TIGR Reptile Database ” (former EMBL Reptile Database)]
<==o Varanus Merrem, 1820 [Hydrosaurus Wagler, 1830; Megalania; Notiosaurus; Odantria] ?L.Eoc. As., Olig.-R. Aust., M.Mioc.-R. As.; L.Mioc.-R. Af., Mioc. Eu. (varaanit; monitor lizards) [46 species] |--+?- V. albigularis sensu Conrad et al., 2008 (valkokurkkuvaraani) | |-- V. griseus (Deset monitor; aavikkovaraani) | `--+-- V. exanthematicus (Steppe monitor; arovaraani) | `-- V. niloticus (Nile monitor; Qamu; niilinvaraani) `--+--+--+--+-- V. rudicollis | | | `-- V. salvator Laurenti, 1768 (juovavaraani) | | `--+-- V. flavescens | | `--+-- V. dumerilii | | `-- V. bengalensis (bengalinvaraani) | `--+--+?- V. bogerti | | |?- V. telenesetes | | |-- V. prasinus | | `--+-- V. beccari | | `-- V. kiethhornei | `--+--+-- V. indicus (Indian monitor; intianvaraani) | | `-- V. melinus | `--+?- V. karlschmidti | |-- V. jobiensis | `--+-- V. doreanus | `-- V. yuwonoi `--+--+--+?- †V. priscus [Megalania prisca; Notiosaurus] sensu Conrad et al., 2008 (megalania) | | `-- V. salvadorii (krokotiilivaraani) | `--+-- V. varius (kirjovaraani) | `-- V. komodoensis Ouwens, 1912 (komodonvaraani; Komodo dragon) `--+--+--+-- V. spenceri | | `-- V. mertensi (mertensinvaraani) | `--+-- V. giganteus | `--+-- V. panoptes (suurvaraani) | `--o V. gouldii (gouldinvaraani) | |-- V. g. gouldii | `-- V. g. flavirufus [V. flavirufus] `--o “Odantria” (kääpiövaraanit; dwarf monitors) |?- V. brevicauda (lyhythäntävaraani) |--+-- V. eremius | `--+-- V. gilleni | `--+--+-- V. kingorum | | `-- V. primodius | `--+-- V. storri | `--+-- V. acanthurus | `-- V. baritji `--+--+-- V. pilbarensis | `-- V. glebopalma `--+--+-- V. glauerti | |?- V. kordensis sensu Conrad et al., 2008 | |?- V. prasinus sensu Conrad et al., 2008 | `--o V. tristis | |-- V. t. tristis | `-- V. t. orientalis `--+-- V. scalaris `--+-- V. timorensis (Timor dwarf monitor; timorinkääpiövaraani) `--+-- V. mitchelli `-- V. semiremex
Untreated Varanus-species:
V. caerulivirens
V. caudolineatus
V. cerambonensis
V. finschi
†V. hofmanni Roger, 1898
V. mabitang Gaulke & Curio, 2001
V. macraei Böhme & Jacobs, 2001
V. pilbarensis
†V. pronini Zerova & Ckhikvadze, 1986
V. rosenbergi
†V. rusingensis Clos, 1995
†V. sivalensis Falconer, 1868
V. spinulosus
V. storri
V. teriae
V. yemenensis (jemeninvaraani, arabianvaraani)
Altenative phylogenies of Varanus:
After Pepin, 199?, and Pianka, 199?
<==o Varanus
|--o (Odantria)
| |?- 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
|
After Conrad, Rieppel & Grande, 2008 (fig. 2 “Adams”)
<==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):
- 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., 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] - 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 TIGR Reptile Database
[former “EMBL Reptile Database”.].
–iNet: http://www.reptile-database.org/