Agamidae

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Agamidae
Temporal range: Cenomanian–recent
MC Siedleragame.jpg
Male Mwanza flat-headed rock agama (Agama mwanzae), in the Serengeti, Tanzania
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Squamata
Suborder: Iguania
Clade: Acrodonta
Family: Agamidae
Gray, 1827 [1]
Subfamilies

6, see text

Agamidae is a family of over 550 species of iguanian lizards indigenous to Africa, Asia, Australia, and a few in Southern Europe. Many species are commonly called dragons or dragon lizards.

Contents

Overview

Phylogenetically, they may be sister to the Iguanidae, and have a similar appearance. Agamids usually have well-developed, strong legs. Their tails cannot be shed and regenerated like those of geckos (and several other families such as skinks), though a certain amount of regeneration is observed in some. [2] [3] Many agamid species are capable of limited change of their colours to regulate their body temperature. [4] In some species, males are more brightly coloured than females, [5] and colours play a part in signaling and reproductive behaviours. [6] Although agamids generally inhabit warm environments, ranging from hot deserts to tropical rainforests, at least one species, the mountain dragon, is found in cooler regions. They are particularly diverse in Australia. [7] [8] [9]

This group of lizards includes some more popularly known, such as the domesticated bearded dragon, Chinese water dragon, and Uromastyx species.

One of the key distinguishing features of the agamids is their teeth, which are borne on the outer rim of their mouths (acrodonts), rather than on the inner side of their jaws (pleurodonts). This feature is shared with the chameleons and the tuatara, but is otherwise unusual among lizards. Agamid lizards are generally diurnal, 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, meaning they are heliothermic. They generally feed on insects and other arthropods (such as spiders), although for some larger species, their diet may include small reptiles or mammals, nestling birds, and flowers or other vegetable matter. [10]

Reproduction

The great majority of agamid species are oviparous. [11] 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 act as a refuge from predators.

Systematics and distribution

Ground agama (Agama aculeata) in Tanzania Lizard - e.JPG
Ground agama (Agama aculeata) in Tanzania
Pondichery fan-throated lizard Sitana ponticeriana from the Agaminae Fan-throated Lizard (Sitana ponticeriana) W IMG 7530.jpg
Pondichéry fan-throated lizard Sitana ponticeriana from the Agaminae
Agamidae in Namibia Agame Canon Lodge Namibia.jpg
Agamidae in Namibia

Very few studies of Agamidae have been conducted. The first comprehensive assessment was by Moody (1980) [12] followed by a more inclusive assessment by Frost and Etheridge (1989). [13] Subsequent studies were based on mitochondrial DNA loci by Macey et al. (2000) [14] and Honda et al. (2000) [15] and also by sampling across the Agamidae by Joger (1991). [16] Few other studies focused on clades within the family, and Agamidae have not been as well investigated as Iguanidae.

The agamids show a curious distribution. They are found over much of the Old World, including continental Africa, Australia, southern Asia, and sparsely in warmer regions of Europe. They are absent from Madagascar and the New World. The distribution is the opposite of that of the iguanids, 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 and pythons. [17]

Subfamilies

Among Agamidae, six subfamilies are generally recognized: [18] [19]

Evolutionary history

The oldest known unambiguous agamid is Protodraco from the mid-Cretaceous (early Cenomanian) aged Burmese amber of Myanmar, dating to around 99 million years ago. It is similar to primitive living Southeast Asian agamids. [20] Gueragama from the Late Cretaceous of Brazil may also be an agamid. [21] Jeddaherdan, a supposed agamid from the Late Cretaceous of Morocco, was later shown to be actually a young subfossil of the living genus Uromastyx. [22]

Predator responses

Body temperature helps determine the physiological state of agamids and affects their predator responses. A positive correlation is seen between a flight response (running speed) and body temperature of various agamid species. [23] 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, play a role in their defensive 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. [24]

Related Research Articles

<span class="mw-page-title-main">Squamata</span> Order of reptiles

Squamata is the largest order of reptiles, comprising lizards and snakes. With over 12,162 species, it is also the second-largest order of extant (living) vertebrates, after the perciform fish. Squamates are distinguished by their skins, which bear horny scales or shields, and must periodically engage in molting. They also possess movable quadrate bones, making possible movement of the upper jaw relative to the neurocranium. This is particularly visible in snakes, which are able to open their mouths very widely to accommodate comparatively large prey. Squamates are the most variably sized living reptiles, ranging from the 16 mm (0.63 in) dwarf gecko to the 6.5 m (21 ft) reticulated python. The now-extinct mosasaurs reached lengths over 14 m (46 ft).

<i>Draco</i> (lizard) Genus of lizards

Draco is a genus of agamid lizards that are also known as flying lizards, flying dragons or gliding lizards. These lizards are capable of gliding flight via membranes that may be extended to create wings (patagia), formed by an enlarged set of ribs. They are arboreal insectivores.

<i>Gonocephalus</i> Genus of lizards

Gonocephalus is a genus of agamid lizards endemic to southeast Asia.

<span class="mw-page-title-main">Thorny devil</span> Species of lizard

The thorny devil, also known commonly as the mountain devil, thorny lizard, thorny dragon, and moloch, is a species of lizard in the family Agamidae. The species is endemic to Australia. It is the sole species in the genus Moloch. It grows up to 21 cm (8.3 in) in total length, with females generally larger than males.

<span class="mw-page-title-main">Frilled lizard</span> Species of reptile

The frilled lizard, also known commonly as the frilled agama, the frillneck lizard, the frill-necked lizard, and the frilled dragon, is a species of lizard in the family Agamidae. The species is native to northern Australia and southern New Guinea and is the only member of the genus Chlamydosaurus. Its common names refer to the large frill around its neck, which usually stays folded against the lizard's body. The frilled lizard grows to 90 cm (35 in) from head to tail tip and can weigh 600 g (1.3 lb). Males are larger and more robust than females. The lizard's body is generally grey, brown, orangish-brown, or black in colour. The frills have red, orange, yellow, or white colours.

<span class="mw-page-title-main">Iguanomorpha</span> Infraorder of lizards

Iguania is an infraorder of squamate reptiles that includes iguanas, chameleons, agamids, and New World lizards like anoles and phrynosomatids. Using morphological features as a guide to evolutionary relationships, the Iguania are believed to form the sister group to the remainder of the Squamata, which comprise nearly 11,000 named species, roughly 2000 of which are iguanians. However, molecular information has placed Iguania well within the Squamata as sister taxa to the Anguimorpha and closely related to snakes. The order has been under debate and revisions after being classified by Charles Lewis Camp in 1923 due to difficulties finding adequate synapomorphic morphological characteristics. Most iguanians are arboreal but there are several terrestrial groups. They usually have primitive fleshy, non-prehensile tongues, although the tongue is highly modified in chameleons. Today they are scattered occurring in Madagascar, the Fiji and Friendly Islands and Western Hemisphere.

<i>Draco dussumieri</i> Species of lizard

Draco dussumieri, also known commonly as the Indian flying lizard, the southern flying lizard, and the Western Ghats flying lizard, is a species of lizard in the family Agamidae. The species is capable of gliding from tree to tree. It is found principally in the Western Ghats and some other hill forests of Southern India. It is almost completely arboreal, found on trees in forests and adjoining palm plantations where it climbs trees to forage for insects and glides to adjoining trees by expanding the patagium, loose skin on the sides of the body which is supported by elongated ribs to act as wings. The skin on the sides of the neck is also extended to the sides using the hyoid bones of the tongue as support. During the breeding season males maintain small territories which they defend from other males while courting females. The male has a more colourful patagium than the female, and it prominently extends its yellow dewlap forward in display. Although living almost its entire life in trees, the female descends to the ground to lay eggs in soil. This is the species with the westernmost distribution within the genus Draco, the majority of species occurring in Southeast Asia.

<i>Saara hardwickii</i> Species of lizard

Saara hardwickii, commonly known as Hardwicke's spiny-tailed lizard or the Indian spiny-tailed lizard is a species of lizard in the family Agamidae. The species is found in patches across the Thar Desert, Kutch, and surrounding arid zones in India and Pakistan. It is mainly herbivorous and lives in numbers in some areas. Since it is found in loose clusters it often attracts predators such as raptors. It is also hunted by local peoples in the belief that the fat extracted from it is an aphrodisiac.

<i>Draco indochinensis</i> Species of lizard

Draco indochinensis, also known as the Indochinese flying lizard or Indochinese gliding lizard, is a species of agamid lizard endemic to South-east Asia.

<i>Ctenophorus pictus</i> Species of lizard

Ctenophorus pictus, commonly known as the painted ground-dragon or painted dragon, is a species of lizard from the family Agamidae. It is endemic to the drier areas of southern and central Australia.

<i>Barbaturex</i> Extinct genus of lizards

Barbaturex is an extinct genus of giant herbivorous iguanian lizards from Pondaung Formation, the Eocene of Myanmar. It is represented by a single species, Barbaturex morrisoni, which is known from several partial dentaries and a fused pair of frontals, two bones that form part of the top of the skull. Based on the size of these bones, Barbaturex morrisoni is estimated to have been about 1 metre (3.3 ft) from snout to vent, and possibly up to 6 feet (1.8 m) including the tail. Barbaturex morrisoni was named after The Doors frontman Jim Morrison, a play on his epithet "The Lizard King". The genus's name is a portmanteau of the Latin words Barbatus and rex, meaning "bearded king", in reference to ridges along the mandible and the lizard's large size.

<span class="mw-page-title-main">Chamaeleontiformes</span> Clade of lizards

Chamaeleontiformes is a hypothesized clade of iguanian lizards defined as all taxa sharing a more recent common ancestor with Chamaeleo chamaeleon than with Hoplocercus spinosus, Polychrus marmoratus, or Iguana iguana. It was named by paleontologist Jack Conrad in 2008 to describe a clade recovered in his phylogenetic analysis that included the extinct genus Isodontosaurus, the extinct family Priscagamidae, and the living clade Acrodonta, which includes agamids and chameleons. It is a stem-based taxon and one of two major clades within Iguania, the other being Pleurodonta. Below is a cladogram from Daza et al. (2012) showing this phylogeny:

Gueragama is an extinct genus of iguanian lizard from the Cretaceous of Brazil. It belongs to a group of iguanians called Acrodonta, whose living members include chameleons and agamids and are currently restricted to the Old World. Gueragama is the only acrodont known from South America, providing evidence that the group once ranged across much of Gondwana and only became restricted to the Old World after the supercontinent broke apart. The type species, Gueragama sulamericana, was named in 2015 on the basis of an isolated lower jaw from Goio-Erê Formation in the Bauru Basin, which was deposited in a desert environment. Although description shows age of Formation at Turonian to Campanian, later study proposes age around Aptian to Albian instead. Unlike modern acrodontans, whose teeth implant on the margins of the jaws, Gueragama has teeth that implant along the inner surface of the lower jaw, a feature common in most non-acrodontan lizards and characteristic of the second major group of iguanians, Pleurodonta. The non-acrodont dentition of Gueragama is evidence of its basal position within Acrodonta, and is shared with the taxon of Late Cretaceous acrodontan relatives, the family Priscagamidae. In 2022, it is considered that is unlikely to be acrodontan, more likely to be scincoids or teiids instead.

Bharatagama is an extinct genus of lepidosaur from the Early Jurassic of India. It has been suggested to be one of the oldest known lizards and the oldest known iguanian. The type and only species is Bharatagama rebbanensis, named in 2002. Over one hundred fossils of Bharatagama have been found in the Kota Formation, which outcrops in the Pranhita–Godavari Basin and dates back to about 190 million years ago (Ma). Despite its abundance, Bharatagama is known only from isolated jaw bones mixed together in microvertebrate assemblages with equally fragmentary remains of fish, sphenodontians, dinosaurs, crocodylomorphs, and mammals. These fossils represent all stages of development, from hatchlings to adults. The total length of the skull in adult specimens is estimated to have been about 15 millimetres (0.59 in). Later analysis suggested that the taxon might be a member of Rhynchocephalia.

<i>Diploderma swinhonis</i> Species of lizard

Diploderma swinhonis, also known commonly as the Taiwan japalure, Swinhoe's japalure, Swinhoe's lizard, and Swinhoe's tree lizard, is a species of lizard in the family Agamidae. The species is native to Taiwan. It is considered an invasive alien species in Japan after likely being transported from Taiwan by humans. A foraging ambush predator, this lizard preys primarily on arthropods and thus remains at the bottom of forests perched on trees where sunlight is present. D. swinhonis is not a major threat to humans and is able to adapt to a variety of habitats, including urban environments. The male D. swinhonis is physically distinct from the female, with its body size being much larger and having a yellow stripe. This species sexually reproduces on a seasonal basis and hibernates during the winter time.

<span class="mw-page-title-main">Acrodonta (lizard)</span> Subclade of lizards

Acrodonta are a subclade of iguanian squamates consisting almost entirely of Old World taxa. Extant representation include the families Chamaeleonidae (chameleons) and Agamidae, with at least over 500 species described. A fossil genus, Gueragama, was found in Brazil, making it the only known American representative of the group.

<i>Lophosaurus</i> Genus of lizards

Lophosaurus is a genus of arboreal agamid lizards from Australia and Melanesia.

<i>Ctenophorus caudicinctus</i> Species of lizard

Ctenophorus caudicinctus, commonly known as the ring-tailed dragon or ring-tailed bicycle-dragon is a native species of agamid lizard occurring in rocky ranges and outcrops of Australia. Ctenophorus caudicinctus is most commonly found in the Pilbara region and offshore islands of Western Australia. The ctenophorus has 28 known species in the northern, southern, and western parts of Australia. It is recognized to be the most speciose group of Australian agamids.

Ctenophorus mckenziei, more commonly known as the dwarf-bicycle dragon, is a species of endemic Australian lizard within the family Agamidae and genus Ctenophorus. Originally identified as the agamid Amphibolurus mckenziei, the lizard had been identified within the regions of Western Australia and South Australia in which it occupied the shrubbery and woodland areas as its habitat. It was subsequently transferred to the genus Ctenophorus along with other Agamid species in which it shared similar morphology and characteristics. The name mckenziei is in reference to Norman Leslie Mckenzie, who was a zoologist and discovered the existence of the lizard. Listed on the IUCN red list page, threats to its population numbers are evaluated as least concern; however, their numbers are threatened by habitat loss, climate change, and feral predators.

<i>Ctenophorus parviceps</i> Species of lizard

Ctenophorus parviceps, commonly known as the Gnaraloo heath dragon or northwestern heath dragon is a species of agamid lizard occurring in pale coastal sands and shell grit with open heaths and beach spinifex, between the North West Cape and Carnarvon, Western Australia and on Bernier Island. The Gnaraloo heath dragon is a lizard that can be found along the coast of Western Australia between Exmouth Gulf and Shark Bay, and is also known as the northwestern heath dragon. It is native to Australia and usually inhabits sandy coastal dunes. The species' longevity is 3–50 years and its population density is extremely low. The Gnaraloo heath dragon is a member of the Agamidae family, which contains 15 genera. The lizard is under the Ctenophorus genus which has up to 33 species. This genus shows the most morphological and ecological diversity out of the three large agamid genera. 83% of the lizards in this genus lack a crest, while 17% possess crests. They are smaller than most agamids but do have relatively large heads. The Gnaraloo heath dragon can be differentiated from related species by a series of spines on the tail's base, a pale-grey brown broad vertebral band along its back, and hour-glass bars extending upwards to meet the pale vertebral band. It is usually 45mm in terms of length, measuring from snout to vent.

References

  1. ↑ "Agamidae". Dahms Tierleben (in German).
  2. ↑ Thompson, M. B. (1993). "Estimate of the population structure of the e[a]stern water dragon, Physignathus lesueurii (Reptilia: Agamidae), along riverside habitat". Wildlife Research. 20 (5): 613–9. doi:10.1071/WR9930613.
  3. ↑ Ananjeva, Natalia B.; Bryan L. Stuart (2001). "The Agamid lizard Ptyctolaemus phuwtilmensis Manthey and Nabhitabhata, 1991 from Thailand and Laos represents a new genus". Russian Journal of Herpetology. 8 (3). Folium Publishing Company: 165–170.
  4. ↑ de Velasco, Jesus Barraza; Glenn J. Tattersall (September 2008). "The influence of hypoxia on the thermal sensitivity of skin colouration in the bearded dragon, Pogona vitticeps". Journal of Comparative Physiology B. 178 (7): 867–875. doi:10.1007/s00360-008-0274-8. PMID   18491114. S2CID   13413178.
  5. ↑ Cuervo, J.J.; R. Shine (10 July 2007). "Hues of a dragon's belly: morphological correlates of ventral coloration in water dragons". Journal of Zoology. 273 (3): 298–304. doi:10.1111/j.1469-7998.2007.00328.x.
  6. ↑ LeBas, Natasha R.; N. Justin Marshall (2000). "The role of colour in signaling and male choice in the agamid lizard Ctenophorus ornatus". Proceedings of the Royal Society of London B. 267 (1442): 445–452. doi:10.1098/rspb.2000.1020. PMC   1690562 . PMID   10737400.
  7. ↑ Gray, JA; Sherratt, E; Hutchinson, MN; Jones, MEH (2019). "Changes in ontogenetic patterns facilitate diversification in skull shape of Australian agamid lizards". BMC Evolutionary Biology. 19 (1): 7. Bibcode:2019BMCEE..19....7G. doi: 10.1186/s12862-018-1335-6 . PMC   6325775 . PMID   30621580.
  8. ↑ Gray JA, Hutchinson MN, Jones ME (2019). "Exceptional disparity in Australian agamid lizards is a possible result of arrival into vacant niche". The Anatomical Record. 302 (9): 1536–1543. doi: 10.1002/ar.24096 . PMID   30773845.
  9. ↑ Gray JA, Sherratt E, Hutchinson MN, Jones ME (2019). "Evolution of cranial shape in a continental-scale evolutionary radiation of Australian lizards" (PDF). Evolution. 73 (11): 2216–29. doi:10.1111/evo.13851. PMID   31580481. S2CID   203652748.
  10. ↑ Cogger, H. G. (1994). Reptiles and Amphibians of Australia (6th ed.). Port Melbourne, Vic.: Reed. ISBN   978-0-7301-0088-1. OCLC   35576956.
  11. ↑ Bauer, Aaron M. (1998). Cogger, H.G.; Zweifel, R.G. (eds.). Encyclopedia of Reptiles and Amphibians. San Diego: Academic Press. pp. 134–136. ISBN   978-0-12-178560-4.
  12. ↑ Moody, S. M. (1980). Phylogenetic relationships and historical biogeographical relationships of the genera in the family Agamidae (Reptilia: Lacertilia) (PhD). Ann Arbor: University of Michigan. 8017324.
  13. ↑ Frost, Darrel R.; Richard Etheridge (28 September 1989). "A phylogenetic analysis and taxonomy of iguanian lizards (Reptilia: Squamata)". University of Kansas Museum of Natural History Miscellaneous Publications. 81. University of Kansas Museum of Natural History: 1–65. Retrieved 5 January 2012.
  14. ↑ Macey, J. Robert; James A. Schulte II; Allan Larson. (2000). "Evolution and phylogenetic information content of mitochondrial genomic structural features illustrated with acrodont lizards". Systematic Biology. 49 (2): 257–277. doi: 10.1093/sysbio/49.2.257 . PMID   12118408..
  15. ↑ Honda, Masanao; Hidetoshi Ota; Mari Kobayashi; Jarujin Nabhitabhata; Hoi-Sen Yong; Showichi Sengoku; Tsutomu Hikida (2000). "Phylogenetic Relationships of the Family Agamidae (Reptilia: Iguania) Inferred from Mitochondrial DNA Sequences" (PDF). Zoological Science. 1991 (3): 616–622. doi:10.2108/0289-0003(2000)17[527:PROTFA]2.0.CO;2. ISSN   0289-0003. S2CID   53607509.
  16. ↑ Joger, Ulrich (1 August 1991). "A Molecular Phylogeny of Agamid Lizards". University of Kansas Museum of Natural History Miscellaneous Publications. 81 (3). American Society of Ichthyologists and Herpetologists: 616–622. doi:10.2307/1446389. JSTOR   1446389.
  17. ↑ Heads, Michael (2014). "3 Global affinities of Australasian Groups §Indian + Pacific Ocean Groups". Biogeography of Australasia: A Molecular Analysis. Cambridge University Press. p. 119. ISBN   9781107041028.
  18. ↑ Agamidae, UniProt Taxonomy
  19. ↑ B. Ananjeva, Natalia (October 2010). "Asian Agamid lizards (Agamidae, Acrodonta, Sauria, Reptilia): Phylogenetic and taxonomic diversity". Taprobanica: The Journal of Asian Biodiversity. 2 (2): 65. doi: 10.4038/tapro.v2i2.3144 .
  20. ↑ Wagner, Philipp; Stanley, Edward L.; Daza, Juan D.; Bauer, Aaron M. (August 2021). "A new agamid lizard in mid-Cretaceous amber from northern Myanmar". Cretaceous Research. 124: 104813. Bibcode:2021CrRes.12404813W. doi:10.1016/j.cretres.2021.104813. S2CID   233704307.
  21. ↑ Apesteguía, Sebastián; Daza, Juan D.; Simões, Tiago R.; Rage, Jean Claude (September 2016). "The first iguanian lizard from the Mesozoic of Africa". Royal Society Open Science. 3 (9): 160462. Bibcode:2016RSOS....360462A. doi:10.1098/rsos.160462. ISSN   2054-5703. PMC   5043327 . PMID   27703708.
  22. ↑ Vullo, Romain; Bailon, Salvador; Dauphin, Yannicke; Monchot, Hervé; Allain, Ronan (November 2022). "A reappraisal of Jeddaherdan aleadonta (Squamata: Acrodonta), the purported oldest iguanian lizard from Africa". Cretaceous Research. 143: 105412. doi: 10.1016/j.cretres.2022.105412 . S2CID   253349389.
  23. ↑ Hertz, Paul E.; Huey, Raymond B.; Nevo, Eviatar (1 August 1982). "Fight versus flight: Body temperature influences defensive responses of lizards". Animal Behaviour. 30 (3): 676–679. doi:10.1016/S0003-3472(82)80137-1. ISSN   0003-3472. S2CID   53182102.
  24. ↑ Shine, Richard (May 1990). "Function and evolution of the frill of the Frillneck Lizard, Chlamydosaurus Kingii (sauria: Agamidae)". Biological Journal of the Linnean Society. 40 (1): 11–20. doi:10.1111/j.1095-8312.1990.tb00531.x.
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