# Agamidae

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{{Short description|Family of lizards}}
{{Use dmy dates|date=January 2020}}
{{confused|Dragon}}
{{Automatic taxobox
| fossil_range = {{fossil range|Cenomanian|recent}}
| image = MC_Siedleragame.jpg
| image_caption = Male [Mwanza flat-headed rock agama](/source/Mwanza_flat-headed_rock_agama) (''Agama mwanzae''), in the [Serengeti](/source/Serengeti), [Tanzania](/source/Tanzania)
| taxon = Agamidae
| authority = [Gray](/source/John_Edward_Gray), 1827<ref>{{cite web |title=Agamidae |lang=de |website=Dahms Tierleben |url=https://www.dahmstierleben.de/systematik/Squamata/agamidae }}</ref>
| subdivision_ranks = Subfamilies
| subdivision = 6, see text
}}

'''Agamidae''' is a [family](/source/Family_(taxonomy)) containing 582 species in 64 genera of [iguania](/source/iguania)n [lizard](/source/lizard)s indigenous to Africa, Asia, Australia, and a few locations in Southern Europe. Many species are commonly called '''dragons''' or '''dragon lizards'''.

==Overview==
[Phylogenetically](/source/phylogenetics), they may be sister to the [Iguanidae](/source/Iguanidae), and have similar appearances. Agamids usually have well-developed, strong legs. Their tails cannot be shed and regenerated like those of [gecko](/source/gecko)s (and several other families such as [skink](/source/skink)s), though a certain amount of regeneration is observed in some.<ref>{{cite journal| last = Thompson | first = M. B.| title = Estimate of the population structure of the e[a]stern water dragon, ''Physignathus lesueurii'' (Reptilia: Agamidae), along riverside habitat | journal = Wildlife Research | volume = 20| issue = 5| pages = 613–9| year =1993 | doi = 10.1071/WR9930613 | bibcode = 1993WildR..20..613T}}</ref><ref>{{Cite journal|last=Ananjeva |first=Natalia B. |author2=Bryan L. Stuart |title=The Agamid lizard ''Ptyctolaemus phuwtilmensis'' Manthey and Nabhitabhata, 1991 from Thailand and Laos represents a new genus |journal=Russian Journal of Herpetology |volume=8 |issue=3 |pages=165–170 |publisher=Folium Publishing Company |year=2001 |url=http://rjh.folium.ru/index.php/rjh/article/view/580 }}</ref> Many agamid species are capable of limited change of their colours to regulate their body temperature.<ref>{{Cite journal| last = de Velasco | first = Jesus Barraza |author2= Glenn J. Tattersall | title = The influence of hypoxia on the thermal sensitivity of skin colouration in the bearded dragon, ''Pogona vitticeps'' | journal = Journal of Comparative Physiology B | volume = 178 | issue = 7| pages =867–875 |date = September 2008| doi = 10.1007/s00360-008-0274-8 | pmid = 18491114 | s2cid = 13413178 }}</ref> In some species, males are more brightly coloured than females,<ref>{{Cite journal| doi = 10.1111/j.1469-7998.2007.00328.x| last = Cuervo | first = J.J. |author2= R. Shine | title = Hues of a dragon's belly: morphological correlates of ventral coloration in water dragons | journal = Journal of Zoology | volume = 273| issue = 3 | pages = 298–304 | date = 10 July 2007 }}</ref> and colours play a part in signaling and reproductive behaviours.<ref>{{Cite journal | doi = 10.1098/rspb.2000.1020 | pmid = 10737400 | pmc = 1690562 | last = LeBas | first = Natasha R. | author2 = N. Justin Marshall | title = The role of colour in signaling and male choice in the agamid lizard ''Ctenophorus ornatus'' | journal = Proceedings of the Royal Society of London B | volume = 267 | issue = 1442 | pages = 445–452 | year = 2000 }}</ref> Although agamids generally inhabit warm environments, ranging from hot [desert](/source/desert)s to tropical [rainforest](/source/rainforest)s, at least one species, the [mountain dragon](/source/mountain_dragon), is found in cooler regions. They are particularly diverse in Australia.<ref>{{Cite journal |last1=Gray |first1=JA|last2=Sherratt |first2=E|last3=Hutchinson |first3=MN|last4=Jones |first4=MEH | year = 2019 | title = Changes in ontogenetic patterns facilitate diversification in skull shape of Australian agamid lizards | journal = BMC Evolutionary Biology | volume = 19 | issue = 1 | page = 7 | doi = 10.1186/s12862-018-1335-6 | pmid = 30621580 | pmc = 6325775 |doi-access=free |bibcode=2019BMCEE..19....7G }}</ref><ref>{{Cite journal | vauthors = Gray JA, Hutchinson MN, Jones ME  | title = Exceptional disparity in Australian agamid lizards is a possible result of arrival into vacant niche | journal =  The Anatomical Record | volume = 302 | issue = 9 | year = 2019 | pages = 1536–1543 | doi = 10.1002/ar.24096 | pmid = 30773845 | doi-access = free }}</ref><ref>{{Cite journal | vauthors = Gray JA, Sherratt E, Hutchinson MN, Jones ME  | title = Evolution of cranial shape in a continental-scale evolutionary radiation of Australian lizards | journal = Evolution | year = 2019 | volume = 73 | issue = 11 | pages = 2216–29 | doi= 10.1111/evo.13851 | pmid = 31580481 | s2cid = 203652748 | url = https://discovery.ucl.ac.uk/id/eprint/10088393/1/DRAGONS_submission1.pdf }}</ref>

This group of lizards includes some more popularly known, such as the domesticated [bearded dragon](/source/bearded_dragon), [Chinese water dragon](/source/Chinese_water_dragon), and ''[Uromastyx](/source/Uromastyx)'' species. 

Members of the genus ''[Draco](/source/Draco_(lizard))'' have modified ribs that form a wing used to glide through their forest habitat.<ref name=":2">{{Cite journal|last=Dehling|first=J. Maximilian|author-link=species:J. Maximilian Dehling|date=2017-12-13|title=How lizards fly: A novel type of wing in animals|journal=PLOS ONE|language=en|volume=12|issue=12|article-number=e0189573|doi=10.1371/journal.pone.0189573|pmc=5728497|pmid=29236777|bibcode=2017PLoSO..1289573D|doi-access=free}}</ref>
[[File:Draco takeoff.gif|thumb|Member of the agamid genus ''[Draco](/source/Draco_(lizard))'' engaging in gliding flight]]
One of the key distinguishing features of the agamids is their teeth, which are borne on the outer rim of their mouths ([acrodont](/source/acrodont)s), rather than on the inner side of their jaws ([pleurodont](/source/pleurodont)s). This feature is shared with the [chameleon](/source/chameleon)s and the [tuatara](/source/tuatara), but is otherwise unusual among lizards. Agamid lizards are generally [diurnal](/source/Diurnality), with good vision, and include a number of arboreal species, in addition to ground and rock dwellers. Most need to bask in the sun to maintain elevated body temperatures (heliothermic). They generally feed on insects and other [arthropod](/source/arthropod)s (such as spiders), although for some larger species, their diet may include small reptiles or mammals, nestling birds, and flowers or other vegetable matter.<ref>{{Cite book |author=Cogger, H. G. |year=1994 |title=Reptiles and Amphibians of Australia |edition=6th |location=Port Melbourne, Vic. |publisher=Reed |isbn=978-0-7301-0088-1 |oclc=35576956}}</ref>

== Reproduction ==
A great many agamid species are [oviparous](/source/oviparous).<ref name="EoR">{{cite book |editor1=Cogger, H.G. |editor2=Zweifel, R.G. |author= Bauer, Aaron M.|year=1998|title=Encyclopedia of Reptiles and Amphibians|publisher= Academic Press|location=San Diego|pages= 134–136|isbn= 978-0-12-178560-4}}</ref> The eggs are mostly found in damp soil or rotting logs to retain enough moisture during the incubation period. The clutch size varies from four to 10 eggs for most species, and incubation period lasts around 6–8 weeks. Specifically in the Leiolepidinae subfamily of agamids, all species use a burrowing system that reaches moist soil, where eggs are deposited in late spring/early summer or at the beginning of the dry season. The Leiolepidinae burrow system is also used for daily or seasonal retreats, as it allows them to regulate their body temperature or gain refuge from predators.

==Systematics and distribution==
[[File:Lizard - e.JPG|thumb|[Ground agama](/source/Ground_agama) (''Agama aculeata'') in Tanzania]]
[[File:Fan-throated Lizard (Sitana ponticeriana) W IMG 7530.jpg|thumb|[Pondichéry fan-throated lizard](/source/Pondich%C3%A9ry_fan-throated_lizard) (''Sitana ponticeriana'') from the [Agaminae](/source/Agaminae)]]
[[File:Agame Canon Lodge Namibia.jpg|thumb|Agamidae in [Namibia](/source/Namibia)]]
Very few studies of the Agamidae have been conducted. The first comprehensive assessment was by Moody (1980),<ref>{{cite thesis | type = PhD |last = Moody |first=S. M. | year = 1980 | title = Phylogenetic relationships and historical biogeographical relationships of the genera in the family Agamidae (Reptilia: Lacertilia) | publisher = University of Michigan | location = Ann Arbor |id=8017324 |url=https://www.proquest.com/openview/f6c0a7c11b44e60d27269b0226315a31/1 }}</ref> followed by a more inclusive assessment by Frost and Etheridge (1989).<ref>{{Cite journal | last = Frost | first = Darrel R. |author2= Richard Etheridge | title = A phylogenetic analysis and taxonomy of iguanian lizards (Reptilia: Squamata) | journal = University of Kansas Museum of Natural History Miscellaneous Publications | volume = 81 | pages = 1–65 | publisher = University of Kansas Museum of Natural History | date = 28 September 1989 | url = https://www.biodiversitylibrary.org/item/55046 | access-date = 5 January 2012}}</ref> Subsequent studies were based on [mitochondrial DNA](/source/mitochondrial_DNA) loci by Macey ''et al.'' (2000)<ref>{{Cite journal|last=Macey |first=J. Robert |author2=James A. Schulte II |author3=Allan Larson. |title=Evolution and phylogenetic information content of mitochondrial genomic structural features illustrated with acrodont lizards |journal=Systematic Biology |volume=49 |issue=2 |pages=257–277 |year=2000 |doi=10.1093/sysbio/49.2.257 |pmid=12118408 |doi-access=free }}.</ref> and Honda ''et al.'' (2000)<ref>{{Cite journal | last = Honda | first = Masanao |author2=Hidetoshi Ota|author3=Mari Kobayashi|author4=Jarujin Nabhitabhata|author5=Hoi-Sen Yong|author6=Showichi Sengoku|author7= Tsutomu Hikida | title = Phylogenetic Relationships of the Family Agamidae (Reptilia: Iguania) Inferred from Mitochondrial DNA Sequences | journal =  Zoological Science| volume = 1991 | number= 3 | pages = 616–622 | year = 2000 | doi = 10.2108/0289-0003(2000)17[527:PROTFA]2.0.CO;2 | s2cid = 53607509 | url = http://repository.kulib.kyoto-u.ac.jp/dspace/bitstream/2433/57223/1/K093.pdf | issn = 0289-0003 }}</ref> and also by sampling across the Agamidae by Joger (1991).<ref>{{Cite journal | last = Joger | first = Ulrich | title = A Molecular Phylogeny of Agamid Lizards | journal = University of Kansas Museum of Natural History Miscellaneous Publications | volume = 81 | issue = 3 | pages = 616–622 | publisher = American Society of Ichthyologists and Herpetologists | date = 1 August 1991 | doi = 10.2307/1446389 | jstor=1446389}}</ref> Few other studies focused on clades within the family, and the Agamidae have not been as well investigated as the Iguanidae.

The agamids show a curious distribution. They are found over much of the [Old World](/source/Old_World), including continental Africa, Australia, southern Asia, and sparsely in warmer regions of Europe. They are absent from [Madagascar](/source/Madagascar) and the [New World](/source/New_World). The distribution is the opposite of that of the [iguanids](/source/iguanidae), which are found in just these areas, but absent in areas where agamids are found. A similar faunal divide is found in between the [boas](/source/Boidae) and [pythons](/source/Pythonidae).<ref>{{Cite book |chapter=3 Global affinities of Australasian Groups §Indian + Pacific Ocean Groups |title=Biogeography of Australasia: A Molecular Analysis|last=Heads|first=Michael|publisher=Cambridge University Press|year=2014|isbn=978-1-107-04102-8|pages=119 }}</ref>

===Further classification===

Among the Agamidae, six [subfamilies](/source/subfamilies) are generally recognized:<ref>[https://www.uniprot.org/taxonomy/81953 Agamidae], UniProt Taxonomy</ref><ref>{{Cite journal|title=Asian Agamid lizards (Agamidae, Acrodonta, Sauria, Reptilia): Phylogenetic and taxonomic diversity|journal = Taprobanica: The Journal of Asian Biodiversity|volume = 2|issue = 2|page = 65|last=B. Ananjeva|first=Natalia|date=October 2010|doi=10.4038/tapro.v2i2.3144|doi-access=free}}</ref>

* [Agaminae](/source/Agaminae) (Africa, Europe, and South Asia)
* [Amphibolurinae](/source/Amphibolurinae) (Australia and New Guinea, one species in Southeast Asia)
* [Draconinae](/source/Draconinae) (South and Southeast Asia)
* [Hydrosaurinae](/source/Hydrosaurinae) (''Hydrosaurus'', Papua New Guinea, the Philippines, and Indonesia)
* [Leiolepidinae](/source/Leiolepidinae) (''Leiolepis'', Southeast Asia)
* [Uromastycinae](/source/Uromastycinae) (''Saara'' and ''Uromastyx'', Africa and South Asia)
These can be further split into these 64 genera:

* ''[Acanthocercus](/source/Acanthocercus)'' ''(15 species)''
* ''[Acanthosaura](/source/Acanthosaura)'' ''(20 species)''
* ''[Agama](/source/Agama_(lizard))'' ''(47 species)''
* ''[Agasthyagama](/source/Agasthyagama)'' ''(2 species)''
* ''[Amphibolurus](/source/Amphibolurus)'' ''(4 species)''
* ''[Aphaniotis](/source/Aphaniotis)'' ''(3 species)''
* ''[Bronchocela](/source/Bronchocela)'' ''(15 species)''
* ''[Bufoniceps](/source/Bufoniceps_laungwalaensis)'' ''(1 species)''
* ''[Calotes](/source/Calotes)'' ''(29 species)''
* ''[Ceratophora](/source/Ceratophora)'' ''(6 species)''
* ''[Chelosania](/source/Chelosania)'' ''(1 species)''
* ''[Chlamydosaurus](/source/Frilled_lizard)'' ''(1 species)''
* ''[Complicitus](/source/Blackthroated_bloodsucker)'' ''(1 species)''
* ''[Cophotis](/source/Cophotis)'' ''(2 species)''
* ''[Coryphophylax](/source/Coryphophylax) (2 species)''
* ''[Cristidorsa](/source/Cristidorsa)'' ''(2 species)''
* ''[Cryptagama](/source/Cryptagama)'' ''(1 species)''
* ''[Ctenophorus](/source/Ctenophorus)'' ''(38 species)''
* ''[Dendragama](/source/Dendragama)'' ''(4 species)''
* ''[Diploderma](/source/Diploderma)'' ''(47 species)''
* ''[Diporiphora](/source/Diporiphora)'' ''(28 species)''
* ''[Draco](/source/Draco_(lizard))'' ''(41 species)''
* ''[Gonocephalus](/source/Gonocephalus)'' ''(17 species)''
* ''[Gowidon](/source/Gowidon)'' ''(1 species)''
* ''[Harpesaurus](/source/Harpesaurus)'' ''(6 species)''
* ''[Hydrosaurus](/source/Hydrosaurus)'' ''(5 species)''
* ''[Hypsicalotes](/source/Hypsicalotes)'' ''(1 species)''
* ''[Hypsilurus](/source/Hypsilurus)'' ''(18 species)''
* ''[Intellagama](/source/Australian_water_dragon)'' ''(1 species)''
* ''[Japalura](/source/Japalura)'' ''(8 species)''
* ''[Laodracon](/source/species%3ALaodracon) (1 species)''
* ''[Laudakia](/source/Laudakia)'' ''(13 species)''
* ''[Leiolepis](/source/Leiolepis)'' ''(10 species)''
* ''[Lophocalotes](/source/Lophocalotes)'' ''(2 species)''
* ''[Lophognathus](/source/Lophognathus)'' ''(2 species)''
* ''[Lophosaurus](/source/Lophosaurus)'' ''(3 species)''
* ''[Lyriocephalus](/source/Lyriocephalus)'' ''(1 species)''
* ''[Malayodracon](/source/Malayodracon)'' ''(1 species)''
* ''[Mantheyus](/source/Phuwua_rock_agama)'' ''(1 species)''
* ''[Microauris](/source/Microauris)'' ''(1 species)''
* ''[Moloch](/source/Thorny_devil)'' ''(1 species)''
* ''[Monilesaurus](/source/Monilesaurus)'' ''(4 species)''
* ''[Otocryptis](/source/Otocryptis)'' ''(2 species)''
* ''[Paralaudakia](/source/Paralaudakia)'' ''(8 species)''
* ''[Pelturagonia](/source/Pelturagonia)'' ''(5 species)''
* ''[Phoxophrys](/source/Phoxophrys)'' ''(1 species)''
* ''[Phrynocephalus](/source/Phrynocephalus)'' ''(36 species)''
* ''[Physignathus](/source/Chinese_water_dragon)'' ''(1 species)''
* ''[Pogona](/source/Pogona)'' ''(6 species)''
* ''[Psammophilus](/source/Psammophilus)'' ''(2 species)''
* ''[Pseudocalotes](/source/Pseudocalotes)'' ''(23 species)''
* ''[Pseudocophotis](/source/Pseudocophotis)'' ''(2 species)''
* ''[Pseudotrapelus](/source/Pseudotrapelus)'' ''(6 species)''
* ''[Ptyctolaemus](/source/Ptyctolaemus)'' ''(3 species)''
* ''[Rankinia](/source/Rankinia)'' ''(1 species)''
* ''[Saara](/source/Saara_(lizard))'' ''(3 species)''
* ''[Salea](/source/Salea)'' ''(2 species)''
* ''[Sarada](/source/Sarada_(lizard))'' ''(3 species)''
* ''[Sitana](/source/Sitana)'' ''(15 species)''
* ''[Trapelus](/source/Trapelus)'' ''(13 species)''
* ''[Tropicagama](/source/Tropicagama)'' ''(1 species)''
* ''[Tympanocryptis](/source/Tympanocryptis)'' ''(23 species)''
* ''[Uromastyx](/source/Uromastyx)'' ''(15 species)''
* ''[Xenagama](/source/Xenagama)'' ''(4 species)''

== Evolutionary history ==
The oldest known unambiguous agamid is ''[Protodraco](/source/Protodraco)'' from the mid-Cretaceous (early [Cenomanian](/source/Cenomanian)) aged [Burmese amber](/source/Burmese_amber) of Myanmar, dating to around 99 million years ago. It is similar to primitive, extant Southeast Asian agamids.<ref>{{Cite journal|last1=Wagner|first1=Philipp|last2=Stanley|first2=Edward L.|last3=Daza|first3=Juan D.|last4=Bauer|first4=Aaron M.|date=August 2021|title=A new agamid lizard in mid-Cretaceous amber from northern Myanmar|url=https://linkinghub.elsevier.com/retrieve/pii/S0195667121000604|journal=Cretaceous Research |volume=124|article-number=104813|doi=10.1016/j.cretres.2021.104813|bibcode=2021CrRes.12404813W |s2cid=233704307|url-access=subscription}}</ref> ''[Gueragama](/source/Gueragama)'' from the Late Cretaceous of Brazil may also be an agamid.<ref>{{Cite journal|last1=Apesteguía|first1=Sebastián|last2=Daza|first2=Juan D.|last3=Simões|first3=Tiago R.|last4=Rage|first4=Jean Claude|date=September 2016|title=The first iguanian lizard from the Mesozoic of Africa|journal=Royal Society Open Science |volume=3|issue=9|article-number=160462|doi=10.1098/rsos.160462|doi-access=free|issn=2054-5703|pmc=5043327|pmid=27703708|bibcode=2016RSOS....360462A}}</ref> ''Jeddaherdan'', a supposed agamid from the Late Cretaceous of Morocco, was later shown to be actually a young [subfossil](/source/subfossil) of the living genus ''Uromastyx''.<ref>{{Cite journal |last1=Vullo |first1=Romain |last2=Bailon |first2=Salvador |last3=Dauphin |first3=Yannicke |last4=Monchot |first4=Hervé |last5=Allain |first5=Ronan |date=November 2022 |title=A reappraisal of Jeddaherdan aleadonta (Squamata: Acrodonta), the purported oldest iguanian lizard from Africa |journal=Cretaceous Research |volume=143 |language=en |article-number=105412 |doi=10.1016/j.cretres.2022.105412|s2cid=253349389 |doi-access=free }}</ref>

==Predator responses==
Body temperature helps determine the [physiological](/source/Physiology) state of agamids and affects their [predator](/source/Predation) responses. A [positive correlation](/source/Correlation) is seen between a flight response (running speed) and body temperature of various agamid species.<ref>{{Cite journal|last1=Hertz|first1=Paul E.|last2=Huey|first2=Raymond B.|last3=Nevo|first3=Eviatar|date=1982-08-01|title=Fight versus flight: Body temperature influences defensive responses of lizards|url=https://www.sciencedirect.com/science/article/pii/S0003347282801371|journal=Animal Behaviour|language=en|volume=30|issue=3|pages=676–679|doi=10.1016/S0003-3472(82)80137-1|s2cid=53182102|issn=0003-3472|url-access=subscription}}</ref> At higher body temperatures, these lizards tend to flee quickly from predators, whereas at lower temperatures, they tend to have a reduced running speed and show an increased fight response, where they are more likely to be aggressive and attack predators.

Certain physical features of some lizards of these species, such as [frilled-neck lizards](/source/Chlamydosaurus), play a role in their [defensive](/source/Defence_mechanism) responses, as well. During the mating season, males tend to display more of their frill, and give fight responses more often. Both males and females display their frills when they are threatened by predators and during social interactions.<ref>{{Cite journal|last=Shine|first=Richard|date=May 1990|title=Function and evolution of the frill of the Frillneck Lizard, Chlamydosaurus Kingii (sauria: Agamidae)|journal=Biological Journal of the Linnean Society|volume=40 |issue=1|pages=11–20 |doi=10.1111/j.1095-8312.1990.tb00531.x}}</ref>

==References==
{{Reflist}}

==External links==
{{Commons category}}
{{Wikispecies}}
* [http://reptile-database.reptarium.cz/search.php?taxon=Agamidae&submit=Search Agamidae] (all species) at [http://www.reptile-database.org/ The Reptile Database]
* {{cite web |title=''Agamidae'' |publisher=Atlas of Living Australia |url=https://bie.ala.org.au/species/urn:lsid:biodiversity.org.au:afd.taxon:b1930792-c83c-4497-a7f6-a28c5b10d9ae}}
* [Laodracon](/source/species%3ALaodracon). Wikispecies.

{{Squamata families}}
{{Taxonbar|from=Q191102}}
{{Authority control}}

Category:Agamidae
Category:Extant Cenomanian first appearances
Category:Lizard families
Category:Taxa named by John Edward Gray

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Adapted from the Wikipedia article [Agamidae](https://en.wikipedia.org/wiki/Agamidae) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Agamidae?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.
