The Carphodactylidae, informally known as the southern padless geckos, are a

family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...

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 throughout the world. They range from . Geckos a ...

lizards Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia altho ...

in the infraorder

Gekkota 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 throughout the world. They range from . Geckos ar ...

. The family consists of 32 described

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...

in 7

genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...

, all of which are

endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...

Australia Australia, officially the Commonwealth of Australia, is a sovereign country comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands. With an area of , Australia is the largest country by ...

. They belong to the superfamily Pygopodoidea (or Diplodactyloidea), an ancient group of east

Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final sta ...

n geckos now only found in

Australasia Australasia is a region that comprises Australia, New Zealand and some neighbouring islands in the Pacific Ocean. The term is used in a number of different contexts, including geopolitically, physiogeographically, philologically, and ecologic ...

. Despite their well-developed limbs, molecular phylogenies have demonstrated that Carphodactylidae is the

sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...

to Pygopodidae, a highly specialized family of legless lizards. Carphodactylids, despite being the most species-poor family of geckos, are still diverse in habits. Many have unusual, specialized tails with reduced rates of

autotomy Autotomy (from the Greek ''auto-'', "self-" and ''tome'', "severing", αὐτοτομία) or self-amputation, is the behaviour whereby an animal sheds or discards one or more of its own appendages, usually as a self-defense mechanism to elude ...

. They lack adhesive toepads and instead cling to bark or substrate with sharply curved claws and a limited array of lamellae. Carphodactylids are relatively large by gecko standards; most are

nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...

and all are

oviparous Oviparous animals are animals that lay their eggs, with little or no other embryonic development within the mother. This is the reproductive method of most fish, amphibians, most reptiles, and all pterosaurs, dinosaurs (including birds), and m ...

, with a typical clutch size of two eggs. Unlike most Australian geckos, species diversity is concentrated in humid forests along the northeastern edge of Australia. Nevertheless, some genera are diverse in arid regions as well.

Genera

The following

are considered members of the Carphodactylidae:, , (2004). "Phylogenetic relationships among gekkotan lizards inferred from c-mos nuclear DNA sequences and a new classification of the Gekkota". ''Biological Journal of the Linnean Society'' 83: 353– 368. doi:10.1111/j.1095-8312.2004.00393.x, (2011). "Systematics and evolution of the Australian knob-tail geckos (''Nephrurus'', Carphodactylidae, Gekkota): plesiomorphic grades and biome shifts through the Miocene". ''Molecular Phylogenetics and Evolution'' 59 (3): 664-674.

Classification

Historical usage (Carphodactylini)

Members of the family now known as Carphodactylidae were first grouped together in

Kluge Kluge (, ) is a German-derived surname. In German, capitalizing, and adding a final to, the adjective (meaning "clever"), creates a noun meaning "clever one". Although the adjective is a feminine form, the noun can be feminine, neuter or masc ...

(1967)’s reconfiguration of gecko systematics. He named a

tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confl ...

, Carphodactylini, which included ''Carphodactylus'', ''Nephrurus'', and ''Phyllurus'' ("padless carphodactylins") as close relatives in a subgroup. The tribe also included '' Pseudothecadactylus'' and all

New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island coun ...

and

New Caledonia ) , anthem = "" , image_map = New Caledonia on the globe (small islands magnified) (Polynesia centered).svg , map_alt = Location of New Caledonia , map_caption = Location of New Caledonia , mapsize = 290px , subdivision_type = Sovereign st ...

n geckos ("padded carphodactylins") in another subgroup. Carphodactylini was diagnosed by a large patch of preanal pores formed by multiple rows, though these pores were noted as vestigial in ''Nephrurus'' and absent in ''Phyllurus''. Under Kluge’s classification scheme, Carphodactylini was the

to Diplodactylini, which contained other Australian legged geckos. Both tribes formed the

subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus Genus ( plural genera ) is a taxonomic rank used in the biological classifica ...

Diplodactylinae The Diplodactylidae are a family in the suborder Gekkota (geckos), with over 150 species in 25 genera. These geckos occur in Australia, New Zealand, and New Caledonia. Diplodactylids are the most ecologically diverse and widespread family of gec ...

within the

Pygopodidae, alongside the legless lizards of the subfamily Pygopodinae. "Padless carphodactylins" (particularly ''Carphodactylus'') were considered the most primitive and generalized members of Carphodactylini, and by extension Diplodactylinae. A morphological phylogenetic analysis by Bauer (1990) generally supported Kluge’s classification scheme, though the ingroup relations of carphodactylins was very different. Summary of Kluge's classification scheme: Results of Bauer, 1990: However, the

monophyly In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic ...

and structure of Carphodactylini (as defined by morphological traits) started to become unclear once molecular phylogenies were developed.

King King is the title given to a male monarch in a variety of contexts. The female equivalent is queen regnant, queen, which title is also given to the queen consort, consort of a king. *In the context of prehistory, antiquity and contempora ...

(1987) found that ''Nephrurus'' and ''Phyllurus'' apparently retained an ancestral

chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins ar ...

structure, but that ''Carphodactylus'', "padded carphodactylins", and several species of ''

Oedura ''Oedura'' is a genus of medium to large geckos, lizards in the family Diplodactylidae. The genus is endemic to Australia. Species in the genus are referred to by the common name velvet geckos. Geographic range Most of the species of ''Oedura ...

'' (a supposed diplodactylin) shared a derived structure, necessitating a major revision of Kluge’s structure. However, there is much debate over whether chromosomes are useful for inferring gecko systematics. Donnellan ''et al''. (1999) estimated gecko relationships using ''C-mos'', a nuclear gene conserved throughout squamates. Their results considered Carphodactylini to be a

paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...

assemblage, with ''

Rhacodactylus ''Rhacodactylus'' is a genus of medium to large geckos of the family Diplodactylidae. All species in this genus are found on the islands that make up New Caledonia. Genus characteristics include long limbs and toes with well-developed lamell ...

'' and '' Pseudothecadactylus'' incrementally closer to diplodactylins (including ''Oedura''). Simplified results of Donnellan ''et al.'', 1999, focusing on Australasian geckos:

Modern usage (Carphodactylidae)

Carphodactylidae, in its present-day usage, was codified by Han ''et al''. (2004). They expanded on the study of Donnellan ''et al''. (1999), and found that Carphodactylini and Diplodactylinae, as previously used, were polyphyletic. "Padless carphodactylins" formed a clade sister to pygopodines, while "padded carphodactylins" formed a grade of geckos basal to diplodactylins. This prompted several renames of major clades. The clade of "padless carphodactylins" was given the new family Carphodactylidae, while the family Diplodactylidae was erected to encompass traditional diplodactylins and "padded carphodactylins". The family Pygopodidae was basically restricted to the legless pygopodines, and subsequent studies would rename the Australasian radiation to Pygopodoidea or Diplodactyloidea. Simplified results of Han ''et al.'', 2004, using new names as defined in the study: Although placing carphodactylids sister to pygopodids was an unusual result, nearly all subsequent molecular phylogenies have supported it. Shared morphological traits are scant, but the two families both have an unusually complex

squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone. In most tetrapods, the squamosal and quadratojugal bones form the cheek series of the skull. The bone forms an ancestral co ...

with its lower end wrapping around the rear branch of the parietal. Oliver & Bauer (2011) looked into carphodactylid ingroup relationships in more depth using an analysis based on ''C-mos'' along with RAG1 (a nuclear gene) and ND2 (a mitochondrial gene). They had difficulty with resolving the base of Carphodactylidae, but strongly supported the erection of a new genus, '' Uvidicolus''.

Skipwith Skipwith is a village and civil parish about north-east of Selby and south-east of York in the Selby District of North Yorkshire, England. Until the 1974 local government reorganisation Skipwith was part of the East Riding of Yorkshire. Ma ...

''et al.'' (2019) found a different result while running phylogenetic analyses on 4268

ultra-conserved element An ultra-conserved element (UCE) was originally defined as a genome segment longer than 200 base pairs (bp) that is absolutely conserved, with no insertions or deletions and 100% identity, between orthologous regions of the human, rat, and mouse ge ...

s (UCEs) in diplodactyloid geckos. Their analysis placed ''Orraya'' as the earliest-diverging carphodactylid, suggesting that the ancestral habit for the family may have been similar to the cryptic leaf-tailed genera (''Orraya, Phyllurus, Saltuarius''). Simplified results of the combined nuclear and mitochondrial analysis of Oliver & Bauer, 2011: Simplified results of Skipwith ''et al.'', 2019:

References

{{Carphodactylidae Geckos Geckos of Australia Lizard families Taxa named by Arnold G. Kluge