Agamidae
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Agamidae is a
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
containing 582 species in 64 genera of iguanian
lizard Lizard is the common name used for all Squamata, squamate reptiles other than snakes (and to a lesser extent amphisbaenians), encompassing over 7,000 species, ranging across all continents except Antarctica, as well as most Island#Oceanic isla ...
s indigenous to Africa, Asia, Australia, and a few locations in Southern Europe. Many species are commonly called dragons or dragon lizards.


Overview

Phylogenetically In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
, they may be sister to the
Iguanidae The Iguanidae is a family of lizards composed of the iguanas, chuckwallas, and their prehistoric relatives, including the widespread green iguana. Taxonomy Iguanidae is thought to be the sister group to the Crotaphytidae, collared lizards (fam ...
, and have similar appearances. Agamids usually have well-developed, strong legs. Their tails cannot be shed and regenerated like those of
gecko Geckos are small, mostly carnivorous lizards that have a wide distribution, found on every continent except Antarctica. Belonging to the infraorder Gekkota, geckos are found in warm climates. They range from . Geckos are unique among lizards ...
s (and several other families such as
skink Skinks are a type of lizard belonging to the family (biology), family Scincidae, a family in the Taxonomic rank, infraorder Scincomorpha. With more than 1,500 described species across 100 different taxonomic genera, the family Scincidae is one o ...
s), though a certain amount of regeneration is observed in some. Many agamid species are capable of limited change of their colours to regulate their body temperature. In some species, males are more brightly coloured than females, and colours play a part in signaling and reproductive behaviours. Although agamids generally inhabit warm environments, ranging from hot
desert A desert is a landscape where little precipitation occurs and, consequently, living conditions create unique biomes and ecosystems. The lack of vegetation exposes the unprotected surface of the ground to denudation. About one-third of the la ...
s to tropical
rainforest Rainforests are forests characterized by a closed and continuous tree Canopy (biology), canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforests can be generally classified as tropi ...
s, at least one species, the mountain dragon, is found in cooler regions. They are particularly diverse in Australia. 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 The tuatara (''Sphenodon punctatus'') is a species of reptile endemic to New Zealand. Despite its close resemblance to lizards, it is actually the only extant member of a distinct lineage, the previously highly diverse order Rhynchocephal ...
, 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 (heliothermic). They generally feed on insects and other
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
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.


Reproduction

A great many agamid species are
oviparous Oviparous animals are animals that reproduce by depositing fertilized zygotes outside the body (i.e., by laying or spawning) in metabolically independent incubation organs known as eggs, which nurture the embryo into moving offsprings kno ...
. 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

Very few studies of the Agamidae have been conducted. The first comprehensive assessment was by Moody (1980), followed by a more inclusive assessment by Frost and Etheridge (1989). Subsequent studies were based on
mitochondrial DNA Mitochondrial DNA (mtDNA and mDNA) is the DNA located in the mitochondrion, mitochondria organelles in a eukaryotic cell that converts chemical energy from food into adenosine triphosphate (ATP). Mitochondrial DNA is a small portion of the D ...
loci by Macey ''et al.'' (2000) and Honda ''et al.'' (2000) and also by sampling across the Agamidae by Joger (1991). 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 The "Old World" () is a term for Afro-Eurasia coined by Europeans after 1493, when they became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia in the Eastern Hemisphere, previously ...
, including continental Africa, Australia, southern Asia, and sparsely in warmer regions of Europe. They are absent from
Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
and the
New World The term "New World" is used to describe the majority of lands of Earth's Western Hemisphere, particularly the Americas, and sometimes Oceania."America." ''The Oxford Companion to the English Language'' (). McArthur, Tom, ed., 1992. New York: ...
. 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.


Further classification

Among the Agamidae, six subfamilies are generally recognized: * Agaminae (Africa, Europe, and South Asia) * Amphibolurinae (Australia and New Guinea, one species in Southeast Asia) * Draconinae (South and Southeast Asia) * Hydrosaurinae (''Hydrosaurus'', Papua New Guinea, the Philippines, and Indonesia) * Leiolepidinae (''Leiolepis'', Southeast Asia) * Uromastycinae (''Saara'' and ''Uromastyx'', Africa and South Asia) These can be further split into these 64 genera: * ''
Acanthocercus ''Acanthocercus'' is a genus of lizards in the Family (biology), family Agamidae. The genus is Endemism, endemic to Africa and the Arabian Peninsula. Species ''Nota bene'': A Binomial nomenclature, binomial authority in parentheses indicates ...
'' ''(15 species)'' * '' Acanthosaura'' ''(20 species)'' * '' Agama'' ''(47 species)'' * '' Agasthyagama'' ''(2 species)'' * '' Amphibolurus'' ''(4 species)'' * '' Aphaniotis'' ''(3 species)'' * '' Bronchocela'' ''(15 species)'' * '' Bufoniceps'' ''(1 species)'' * '' Calotes'' ''(29 species)'' * '' Ceratophora'' ''(6 species)'' * '' Chelosania'' ''(1 species)'' * '' Chlamydosaurus'' ''(1 species)'' * '' Complicitus'' ''(1 species)'' * '' Cophotis'' ''(2 species)'' * '' Coryphophylax (2 species)'' * '' Cristidorsa'' ''(2 species)'' * '' Cryptagama'' ''(1 species)'' * '' Ctenophorus'' ''(38 species)'' * '' Dendragama'' ''(4 species)'' * '' Diploderma'' ''(47 species)'' * '' Diporiphora'' ''(28 species)'' * '' Draco'' ''(41 species)'' * '' Gonocephalus'' ''(17 species)'' * '' Gowidon'' ''(1 species)'' * '' Harpesaurus'' ''(6 species)'' * '' Hydrosaurus'' ''(5 species)'' * '' Hypsicalotes'' ''(1 species)'' * '' Hypsilurus'' ''(18 species)'' * '' Intellagama'' ''(1 species)'' * '' Japalura'' ''(8 species)'' * '' Laodracon (1 species)'' * '' Laudakia'' ''(13 species)'' * '' Leiolepis'' ''(10 species)'' * '' Lophocalotes'' ''(2 species)'' * '' Lophognathus'' ''(2 species)'' * '' Lophosaurus'' ''(3 species)'' * '' Lyriocephalus'' ''(1 species)'' * '' Malayodracon'' ''(1 species)'' * '' Mantheyus'' ''(1 species)'' * '' Microauris'' ''(1 species)'' * '' Moloch'' ''(1 species)'' * '' Monilesaurus'' ''(4 species)'' * '' Otocryptis'' ''(2 species)'' * '' Paralaudakia'' ''(8 species)'' * '' Pelturagonia'' ''(5 species)'' * '' Phoxophrys'' ''(1 species)'' * '' Phrynocephalus'' ''(36 species)'' * '' Physignathus'' ''(1 species)'' * '' Pogona'' ''(6 species)'' * '' Psammophilus'' ''(2 species)'' * '' Pseudocalotes'' ''(23 species)'' * '' Pseudocophotis'' ''(2 species)'' * '' Pseudotrapelus'' ''(6 species)'' * '' Ptyctolaemus'' ''(3 species)'' * '' Rankinia'' ''(1 species)'' * '' Saara'' ''(3 species)'' * '' Salea'' ''(2 species)'' * '' Sarada'' ''(3 species)'' * '' Sitana'' ''(15 species)'' * '' Trapelus'' ''(13 species)'' * '' Tropicagama'' ''(1 species)'' * '' Tympanocryptis'' ''(23 species)'' * '' Uromastyx'' ''(15 species)'' * '' Xenagama'' ''(4 species)''


Evolutionary history

The oldest known unambiguous agamid is '' Protodraco'' from the mid-Cretaceous (early
Cenomanian The Cenomanian is, in the International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...
) aged
Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. Th ...
of Myanmar, dating to around 99 million years ago. It is similar to primitive, extant Southeast Asian agamids. '' Gueragama'' from the Late Cretaceous of Brazil may also be an agamid. ''Jeddaherdan'', a supposed agamid from the Late Cretaceous of Morocco, was later shown to be actually a young subfossil of the living genus ''Uromastyx''.


Predator responses

Body temperature helps determine the
physiological Physiology (; ) is the science, scientific study of function (biology), functions and mechanism (biology), mechanisms in a life, living system. As a branches of science, subdiscipline of biology, physiology focuses on how organisms, organ syst ...
state of agamids and affects their
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
responses. A positive correlation is seen between a flight response (running speed) and body temperature of various agamid species. 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.


References


External links


Agamidae
(all species) a
The Reptile Database
* * Laodracon. Wikispecies. {{Authority control Extant Cenomanian first appearances Lizard families Taxa named by John Edward Gray