varanidae
{{Short description|Family of lizards}}
{{automatic taxobox
| name = Varanids
| fossil_range =
Late Cretaceous – Holocene, {{fossilrange|80|0}}
| image = Komodo dragons (Varanus komodoensis) fighting.jpg
| image_caption = Male Komodo dragons (Varanus komodoensis) fighting, Indonesia
| image2 = Saniwa FMNH.jpg
| image2_caption = Fossil of Saniwa, an extinct varanid known from the Eocene of North America
| taxon = Varanidae
| authority = Merrem, 1820
| subdivision_ranks = Genera
| subdivision = *Varanus
}}
The Varanidae are a family of lizards in the superfamily Varanoidea and order Anguimorpha. The family, a group of carnivorous and frugivorous lizards,{{cite journal | vauthors = Welton LJ, Siler CD, Bennett D, Diesmos A, Duya MR, Dugay R, Rico EL, Van Weerd M, Brown RM | display-authors = 6 | title = A spectacular new Philippine monitor lizard reveals a hidden biogeographic boundary and a novel flagship species for conservation | journal = Biology Letters | volume = 6 | issue = 5 | pages = 654–658 | date = October 2010 | pmid = 20375042 | pmc = 2936141 | doi = 10.1098/rsbl.2010.0119 }} includes the living genus Varanus and a number of extinct genera more closely related to Varanus than to the earless monitor lizard (Lanthanotus).{{cite journal | vauthors = Dong L, Wang YQ, Zhao Q, Vasilyan D, Wang Y, Evans SE | title = A new stem-varanid lizard (Reptilia, Squamata) from the early Eocene of China | journal = Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences | volume = 377 | issue = 1847 | pages = 20210041 | date = March 2022 | pmid = 35125002 | pmc = 8819366 | doi = 10.1098/rstb.2021.0041 }} Varanus includes the Komodo dragon (the largest living lizard), crocodile monitor, savannah monitor, the goannas of Australia and Southeast Asia, and various other species with a similarly distinctive appearance. Their closest living relatives are the earless monitor lizard and Chinese crocodile lizard.{{cite journal | vauthors = Fry BG, Vidal N, Norman JA, Vonk FJ, Scheib H, Ramjan SF, Kuruppu S, Fung K, Hedges SB, Richardson MK, Hodgson WC, Ignjatovic V, Summerhayes R, Kochva E | display-authors = 6 | title = Early evolution of the venom system in lizards and snakes | journal = Nature | volume = 439 | issue = 7076 | pages = 584–588 | date = February 2006 | pmid = 16292255 | doi = 10.1038/nature04328 | first12 = V. | first13 = R. | s2cid = 4386245 | first14 = E. | bibcode = 2006Natur.439..584F | first111 = W.C. }} The oldest members of the family are known from the Late Cretaceous of Mongolia.
Taxonomy
The Varanidae were defined (using morphological characteristics) by Estes, de Queiroz and Gauthier (1988) as the clade containing the most recent common ancestor of Lanthanotus and Varanus and all of its descendants.{{cite book | veditors = Estes RJ, Pregill GK | vauthors = de Queiroz K, Gauthier J |date=1988 |title=Phylogenetic Relationships of the Lizard Families: Essays Commemorating Charles L. Camp |chapter=Phylogenetic Relationships within Squamata |publisher=Stanford University Press |isbn=9780804714358 |oclc=16646258 |page=166}} A similar definition was formulated by Conrad et al. (2008) (also using morphological data), who defined the Varanidae as the clade containing Varanus varius, Lanthanotus borneensis, and all descendants of their last common ancestor.{{cite journal | vauthors = Conrad J |year=2008 |title=Phylogeny and systematics of Squamata (Reptilia) based on morphology |journal=Bulletin of the American Museum of Natural History |doi=10.1206/310.1 |hdl=2246/5915 |volume=310 |pages=1–182|s2cid=85271610 }} Using one of these definitions leads to the inclusion of the earless monitor lizard (L. borneensis) in the family Varanidae.
Lee (1997) created a different definition of the Varanidae, defining them as the clade containing Varanus and all taxa more closely related to Varanus than to Lanthanotus;{{cite journal | pmid=20986455 | date=1946 | author1=WARNER R | title=Pectoral girdles vs. Hyobranchia in the snake genera Liotyphlops and Anomalepis | journal=Science | volume=103 | issue=2686 | pages=720–722 | doi=10.1126/science.103.2686.720 }}{{cite journal | vauthors = Lee MS | title = Molecular evidence and marine snake origins | journal = Biology Letters | volume = 1 | issue = 2 | pages = 227–230 | date = June 2005 | pmid = 17148173 | pmc = 1626205 | doi = 10.1098/rsbl.2004.0282 }} this definition explicitly excludes the earless monitor lizard from the Varanidae. Whether L. borneensis is included in or excluded from the Varanidae depends on the author; for example, Vidal et al. (2012) classify the earless monitor lizard as a member of a separate family Lanthanotidae,{{cite journal | vauthors = Vidal N, Marin J, Sassi J, Battistuzzi FU, Donnellan S, Fitch AJ, Fry BG, Vonk FJ, Rodriguez de la Vega RC, Couloux A, Hedges SB | display-authors = 6 | title = Molecular evidence for an Asian origin of monitor lizards followed by Tertiary dispersals to Africa and Australasia | journal = Biology Letters | volume = 8 | issue = 5 | pages = 853–855 | date = October 2012 | pmid = 22809723 | pmc = 3441001 | doi = 10.1098/rsbl.2012.0460 }} while Gauthier et al. (2012) classify it as a member of Varanidae.{{cite journal | vauthors = Gauthier JA, Kearney M, Maisano JA, Rieppel O, Behlke AD |year=2012 |title=Assembling the Squamate Tree of Life: Perspectives from the Phenotype and the Fossil Record |journal=Bulletin of the Peabody Museum of Natural History |doi=10.3374/014.053.0101 |volume=53 |issue=1 |pages=3–308|s2cid=86355757 }}
=Genera=
:Genera marked with {{extinct}} are extinct
Genera included in Varanidae according to Dong et al., 2022
- {{extinct}}Ovoo Norell, Gao, & Conrad, 2008{{cite journal | vauthors = Conrad JL, Balcarcel AM, Mehling CM | title = Earliest example of a giant monitor lizard (Varanus, Varanidae, Squamata) | journal = PLOS ONE | volume = 7 | issue = 8 | pages = e41767 | year = 2012 | pmid = 22900001 | pmc = 3416840 | doi = 10.1371/journal.pone.0041767 | bibcode = 2012PLoSO...741767C | doi-access = free }} (Mongolia, Late Cretaceous)
- {{extinct}}Aiolosaurus Gao and Norell, 2000 (Mongolia, Late Cretaceous)
- {{extinct}}Cherminotus Borsuk-Bialynicka, 1984 (Mongolia, Late Cretaceous)
- {{extinct}}Saniwides Borsuk-Bialynicka, 1984 (Mongolia, Late Cretaceous)
- {{extinct}}Paravaranus Borsuk-Bialynicka, 1984 (Mongolia, Late Cretaceous)
- {{extinct}}Proplatynotia Borsuk-Bialynicka, 1984 (Mongolia, Late Cretaceous)
- {{extinct}}Telmasaurus Gilmore, 1943 (Mongolia, Late Cretaceous)
- {{extinct}}Saniwa Leidy, 1870 (Europe, North America, Eocene)
- {{extinct}}Archaeovaranus Dong et al., 2022 (China, Eocene)
- Varanus Shaw, 1790
=Phylogeny=
Below is a cladogram from Dong et al. 2022.
{{Clade|{{Clade
|1=†Ovoo gurvel
|2={{Clade
|1={{Clade
|2={{Clade
}} }}
|2={{Clade
|1={{Clade
|2=†Proplatynotia longirostrata
}}
|2={{Clade
|1=†Saniwa ensidens
|2={{Clade
|2=Varanus
}} }} }} }} }}|label1=Varanidae}}
Biology
File:Grays Monitor WWP.jpg (Varanus olivaceus) is a tree-dwelling varanid from the Philippines that primarily feeds on fruit]]
Monitor lizards are reputed to be among the most intelligent lizards. Most species forage widely and have large home ranges,{{cite journal | vauthors = Perry G, Garland Jr T |authorlink2=Theodore Garland, Jr. |year=2002 |title=Lizard home ranges revisited: effects of sex, body size, diet, habitat, and phylogeny |journal=Ecology |volume=83 |pages=1870–1885 |url=http://www.biology.ucr.edu/people/faculty/Garland/PerryGarland2002.pdf |doi=10.1890/0012-9658(2002)083[1870:LHRREO]2.0.CO;2 |issue=7}} and many have high stamina.{{cite journal | vauthors = Clemente CJ, Withers PC, Thompson GG |year=2009 |title=Metabolic rate and endurance capacity in Australian varanid lizards (Squamata; Varanidae; Varanus) |journal=Biological Journal of the Linnean Society |doi=10.1111/j.1095-8312.2009.01207.x |volume=97 |issue=3 |pages=664–676|doi-access=free }} Although most species are carnivorous, three arboreal species in the Philippines (Varanus olivaceus, Varanus mabitang, and Varanus bitatawa) are primarily frugivores.{{cite book | vauthors = Greene HW |date=1986 |title=Diet and Arboreality in the Emerald Monitor, Varanus prasinus, with Comments on the Study of Adaptation |publisher=Field Museum of Natural History |location=Chicago |oclc=14915452 |ol=7155983M}} Among species of living varanids, the limbs show positive allometry, being larger in larger-bodied species, although the feet become smaller as compared with the lengths of the other limb segments.{{cite journal | vauthors = Christian A, Garland Jr T |authorlink2=Theodore Garland, Jr. |year=1996 |title=Scaling of limb proportions in monitor lizards (Squamata: Varanidae) |journal=Journal of Herpetology |doi=10.2307/1565513 |jstor=1565513 |volume=30 |issue=2 |pages=219–230 |url=http://www.biology.ucr.edu/people/faculty/Garland/ChriGa96.pdf}}
Varanids possess unidirectional pulmonary airflow, including air-sacs akin to those of birds.[http://svpow.com/2013/12/11/unidirectional-airflow-in-the-lungs-of-birds-crocs-and-now-monitor-lizards/ Unidirectional Airflow In The Lungs Of Birds, Crocs And Now Monitor Lizards]
See also
References
{{Reflist|30em}}
External links
- [http://www.jcvi.org/reptiles/families/varanidae.php Varanidae]
{{Squamata families}}
{{varanoidea}}
{{Taxonbar|from=Q3082684}}
{{Authority control}}