Ngome Dwarf Chameleon
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''Bradypodion'' (meaning "slow-footed" in
Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...
) is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
chameleon Chameleons or chamaeleons (Family (biology), family Chamaeleonidae) are a distinctive and highly specialized clade of Old World lizards with 200 species described as of June 2015. The members of this Family (biology), family are best known for ...
s in the
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 ...
Chamaeleonidae, collectively called South African dwarf chameleons. All species are found in
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
and most are
endemic Endemism is the state of a species being found only 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 foun ...
to this country, but a few can also be found in
Eswatini Eswatini, formally the Kingdom of Eswatini, also known by its former official names Swaziland and the Kingdom of Swaziland, is a landlocked country in Southern Africa. It is bordered by South Africa on all sides except the northeast, where i ...
,
Lesotho Lesotho, formally the Kingdom of Lesotho and formerly known as Basutoland, is a landlocked country in Southern Africa. Entirely surrounded by South Africa, it is the largest of only three sovereign enclave and exclave, enclaves in the world, t ...
, southernmost
Namibia Namibia, officially the Republic of Namibia, is a country on the west coast of Southern Africa. Its borders include the Atlantic Ocean to the west, Angola and Zambia to the north, Botswana to the east and South Africa to the south; in the no ...
and possibly southernmost
Mozambique Mozambique, officially the Republic of Mozambique, is a country located in Southeast Africa bordered by the Indian Ocean to the east, Tanzania to the north, Malawi and Zambia to the northwest, Zimbabwe to the west, and Eswatini and South Afr ...
. They are quite small chameleons where the different species often can be difficult to separate by appearance, although exact location (each species tends to have a rather small distribution) and the intense breeding colours of males are useful for their identification. They are
arboreal Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally (scansorial), but others are exclusively arboreal. The hab ...
, but some species are mostly found low in the vegetation. ''Bradypodion'' feed on small invertebrates, especially insects. They rapidly reach sexual maturity, typically at an age between half a year and one year, but sometimes up to two years. Adult males are
territorial A territory is an area of land, sea, or space, belonging or connected to a particular country, person, or animal. In international politics, a territory is usually a geographic area which has not been granted the powers of self-government, ...
. It is the only genus of chameleon where females of all species give birth to live young rather than lay eggs (the only other chameleons that give birth to live young are certain ''
Trioceros ''Trioceros'' is a genus of lizards in the family Chamaeleonidae, the chameleons, native to lowlands and highlands in the African mainland, ranging from Ethiopia south to Mozambique and west as far as Ghana. ''Trioceros'' was considered a subgenus ...
'' species). This is likely an adaption to the relatively cold climate of southern Africa in comparison to tropical Africa. Colder temperatures slow the development of eggs laid in the ground; when instead retained inside the body until birth, a female can actively sun bask to increase the temperature. A female can give birth to up to 20 young per time, twice or rarely even three times in a year. Up until the early 2000s, some other chameleons from eastern and central Africa were occasionally placed herein, but they are now placed in ''
Kinyongia ''Kinyongia'' (derived from the group's name in Kiswahili language) is a genus of chameleons found in montane and sub-montane areas in Kenya, Tanzania, Uganda, Rwanda, Burundi, and far eastern DR Congo. They are restricted to forests, woodlands a ...
'' and ''
Nadzikambia ''Nadzikambia'' (derived from their name in Chichewa) is a genus of chameleons. It includes two species of small, plesiomorphic chameleons from the Ruo Gorge forest on Mount Mulanje in Malawi and Mount Mabu in Mozambique. Initially only a sin ...
''.


Species

20 species are currently recognized:


Undescribed species

* Emerald dwarf chameleon, ''Bradypodion'' sp. (Emerald) * ''Bradypodion'' sp. (Groendal) * ''Bradypodion'' sp. (Grootvadersbosch) * ''Bradypodion'' sp. (Jagersbos)


Systematics

Delimitation of ''Bradypodion'' has been controversial for some time. Most species seem readily distinguishable by morphological characteristics, but for some time the genus was used as a
wastebin taxon Wastebasket taxon (also called a wastebin taxon, dustbin taxon or catch-all taxon) is a term used by some taxonomists to refer to a taxon that has the purpose of classifying organisms that do not fit anywhere else. They are typically defined by e ...
for smaller chameleons from
sub-Saharan Africa Sub-Saharan Africa is the area and regions of the continent of Africa that lie south of the Sahara. These include Central Africa, East Africa, Southern Africa, and West Africa. Geopolitically, in addition to the list of sovereign states and ...
with
plesiomorph In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral Phenotypic trait, character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorph ...
ic
hemipenis A hemipenis (: hemipenes) is one of a pair of intromittent organs of male squamates (snakes and lizards). Hemipenes are usually held inverted within the body, and are everted for reproduction via erectile tissue, much like that in the human pe ...
es. Alternatively, many of the present species were reduced to subspecies status. This has since been refuted, but several more species seem recognizable judging from morphological and
mitochondrial A mitochondrion () is an organelle found in the cells of most eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is used ...
16S rRNA 16S ribosomal RNA (or 16Svedberg, S rRNA) is the RNA component of the 30S subunit of a prokaryotic ribosome (SSU rRNA). It binds to the Shine-Dalgarno sequence and provides most of the SSU structure. The genes coding for it are referred to as ...
and
NADH dehydrogenase NADH dehydrogenase is an enzyme that converts nicotinamide adenine dinucleotide (NAD) from its reduced form (NADH) to its oxidized form (NAD+). Members of the NADH dehydrogenase family and analogues are commonly systematically named using the f ...
subunit 2
sequence In mathematics, a sequence is an enumerated collection of objects in which repetitions are allowed and order matters. Like a set, it contains members (also called ''elements'', or ''terms''). The number of elements (possibly infinite) is cal ...
data.Tolley ''et al.'' (2004) The
phylogeny A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or Taxon, taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, M ...
and
biogeography Biogeography is the study of the species distribution, distribution of species and ecosystems in geography, geographic space and through evolutionary history of life, geological time. Organisms and biological community (ecology), communities o ...
of this group is quite consistently resolved. The
Cape dwarf chameleon The Cape dwarf chameleon (''Bradypodion pumilum'') is a chameleon native to the South African province of the Western Cape, where it is restricted to the region around Cape Town. As with most chameleons, its tongue is twice the length of its body ...
and the Knysna dwarf chameleon (and possibly one new species close to it) are basal lineages with unclear relationships; they seem a bit closer to each other than to any other species, but altogether are quite distant. They occur in isolated ranges in coastal
Western Cape The Western Cape ( ; , ) is a provinces of South Africa, province of South Africa, situated on the south-western coast of the country. It is the List of South African provinces by area, fourth largest of the nine provinces with an area of , an ...
and western
Eastern Cape The Eastern Cape ( ; ) is one of the nine provinces of South Africa. Its capital is Bhisho, and its largest city is Gqeberha (Port Elizabeth). Due to its climate and nineteenth-century towns, it is a common location for tourists. It is also kno ...
provinces. Inhabiting a wide range of
habitat In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...
s, they are (for the genus) large, and have brilliant, predominantly green coloration and long tails - just as in many ''
Chamaeleo ''Chamaeleo'' is a genus of chameleons in the Family (biology), family Chamaeleonidae. Most species of the genus ''Chamaeleo'' are found in sub-Saharan Africa, but a few species are also present in northern Africa, southern Europe, and southern ...
''. These characters are
plesiomorph In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral Phenotypic trait, character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorph ...
ic, retained from the genus' ancestor. The remaining species form a well-supported
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
, which in turn can be divided into smaller groups. One consists of forms that radiated on the seawards slopes of the
Drakensberg Mountains The Drakensberg ( Zulu: uKhahlamba, Sotho: Maloti, Afrikaans: Drakensberge) is the eastern portion of the Great Escarpment, which encloses the central Southern African plateau. The Great Escarpment reaches its greatest elevation – within the ...
: the southern Drakensberg dwarf chameleon, the northern
Transvaal dwarf chameleon The Transvaal dwarf chameleon (''Bradypodion transvaalense'') is a chameleon native to South Africa, where it is found in forested areas of Mpumalanga and Limpopo provinces. It is also known as the Wolkberg dwarf chameleon, after the Wolkberg ran ...
, and what appears to be undescribed species from the Ngome Forest on the southeastern slopes. These are also plesiomorphic in ''habitus'' and habits. Another group of taxa occurs from easternmost Eastern Cape to central
KwaZulu-Natal KwaZulu-Natal (, also referred to as KZN) is a Provinces of South Africa, province of South Africa that was created in 1994 when the government merged the Zulu people, Zulu bantustan of KwaZulu ("Place of the Zulu" in Zulu language, Zulu) and ...
provinces, between Gilboa Forest and the
Tugela River The Tugela River (; ) is the largest river in KwaZulu-Natal Province, South Africa. With a total length of , and a drop of 1370 metres in the lower 480 km, it is one of the most important rivers of the country. The river originates in M ...
. These inhabit a wide range of habitat and contain the plesiomorphic Natal Midlands dwarf chameleon from the namesake region, the small black-headed dwarf chameleon which inhabits
fynbos Fynbos (; , ) is a small belt of natural shrubland or heathland vegetation located in the Western Cape and Eastern Cape provinces of South Africa. The area is predominantly coastal and mountainous, with a Mediterranean climate. The fynbos ...
and other low forest on slopes of mainly coastal KwaZulu-Natal, and another probable new species from the Gilboa Forest area. These appear to be a quite recent
radiation In physics, radiation is the emission or transmission of energy in the form of waves or particles through space or a material medium. This includes: * ''electromagnetic radiation'' consisting of photons, such as radio waves, microwaves, infr ...
from a single ancestor, and the group requires more research as regards species limits, and geographical delimitation from the Drakensberg dwarf chameleon. Several largish but short-tailed and cryptic taxa inhabit more
arid Aridity is the condition of geographical regions which make up approximately 43% of total global available land area, characterized by low annual precipitation, increased temperatures, and limited water availability.Perez-Aguilar, L. Y., Plata ...
habitat In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...
s such as
karoo The Karoo ( ; from the Afrikaans borrowing of the South Khoekhoe Khoemana (also known as !Orakobab or Korana) word is a semidesert natural region of South Africa. No exact definition of what constitutes the Karoo is available, so its extent is ...
. The
Karroo The Karoo ( ; from the Afrikaans borrowing of the South Khoekhoe Khoemana (also known as !Orakobab or Korana) word is a semidesert natural region of South Africa. No exact definition of what constitutes the Karoo is available, so its extent is ...
and southern dwarf chameleons seem to have considerable
gene flow In population genetics, gene flow (also known as migration and allele flow) is the transfer of genetic variation, genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent ...
range from
Northern Cape The Northern Cape ( ; ; ) is the largest and most sparsely populated Provinces of South Africa, province of South Africa. It was created in 1994 when the Cape Province was split up. Its capital is Kimberley, South Africa, Kimberley. It includes ...
to coastal Eastern Cape provinces. The small and nearly
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
Smith's dwarf chameleon is close to these; it occurs on the
escarpment An escarpment is a steep slope or long cliff that forms as a result of faulting or erosion and separates two relatively level areas having different elevations. Due to the similarity, the term '' scarp'' may mistakenly be incorrectly used inte ...
inland from
Jeffreys Bay Jeffreys Bay (, nicknamed J-bay) is a town of 27,107 inhabitants as of the 2011 census in the Eastern Cape province of South Africa. It is the seat of the Kouga municipality and is famous amongst surfers as a surf capital due to the right-han ...
. Less closely related is the
Robertson dwarf chameleon The Robertson dwarf chameleon, also known as the Little Karoo dwarf chameleon, (''Bradypodion gutturale'') is a chameleon in the genus ''Bradypodion''. It is found in the dry Fynbos and Renosterveld shrub vegetation, in the centre of the Western ...
, another aridland species which is found in Western Cape province inland from the range of the Cape dwarf chameleon and may be a
cryptic species complex In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
, and an undescribed population from the
Swartberg Mountains The Swartberg mountains (''black mountain'' in English) are a mountain range in the Western Cape province of South Africa. It is composed of two main mountain chains running roughly east–west along the northern edge of the semi-arid Little Kar ...
. The last species with aridland
apomorph In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to ha ...
ies, the Namaqua dwarf chameleon, is quite distant to the others; it occurs in coastal regions from north of the Cape species through
Namaqualand Namaqualand ( Khoikhoi: "Nama-kwa" meaning Nama Khoi people's land) is an arid region of Namibia and South Africa, extending along the west coast over and covering a total area of . It is divided by the lower course of the Orange River int ...
. This is probably still a part of a single radiation which brought about all the aridland taxa, and eventually Smith's dwarf chameleon. The remaining species are all small inhabitants of forested slopes and fynbos, such as the black-headed and Smith's dwarf chameleons. However, as already indicated by the distinctness of these two, their morphologies seem to be a convergent
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
. The
Kentani Centane, or alternatively anglicised Kentane or Kentani because Europeans often cannot easily pronounce the Xhosa click 'C'; is a settlement in Amathole District Municipality in the Eastern Cape province of South Africa. It is situated at app ...
and
Transkei dwarf chameleon The Transkei dwarf chameleon or Pondo dwarf chameleon (''Bradypodion caffer'') is a chameleon endemic to the Eastern Cape Province of South Africa. Reproduction Transkei dwarf chameleon are ovoviviparous. Habitat and conservation Transkei dwarf ...
s from the east coasts of Eastern Cape may or may not be each other's closest relatives. Setaro's dwarf chameleon from northeastern coastal KwaZulu-Natal is not close to these. The Zululand dwarf chameleon from western uThungulu apparently consists of two or more species, one that may be closer to the preceding, and one that might be an early offshoot of the ancestral Drakensberg stock, and which are distinguishable by morphological and mtDNA characteristics.Raw (2001), Tolley ''et al.'' (2004) In conclusion, of the three basic morphotypes found in this genus, one (bright, long-tailed, large) is plesiomorphic, another (large, short-tailed, drab) apparently only evolved once, and the third (the small, slope-inhabiting forms) are convergent in morphology. The ancestors of ''Bradypodion'' thus were mid-sized chameleons with vivid color, which settled the Cape region from roughly north-northwestwards. Due to climate changes with fluctuating
aridity Aridity is the condition of geographical regions which make up approximately 43% of total global available land area, characterized by low annual precipitation, increased temperatures, and limited water availability.Perez-Aguilar, L. Y., Plata ...
, the basal lineages inhabiting humid fynbos in the southwest became isolated from each other and from the animals living around the border region between Northern and Eastern Cape and Free State, and
Lesotho Lesotho, formally the Kingdom of Lesotho and formerly known as Basutoland, is a landlocked country in Southern Africa. Entirely surrounded by South Africa, it is the largest of only three sovereign enclave and exclave, enclaves in the world, t ...
. The aridland habitat fluctuates in extent during climate shifts, and mountainous habitat becomes fragmented or consolidates accordingly. Consequently, the Drakensberg, the ''B. thamnobates''-''B. melanocephalum'', and the aridland group, as well as several coastal lineages, diverged and evolved to their present-day ranges and diversity.


Footnotes


References

* (1998): ''Field Guide to Snakes and Other Reptiles of Southern Africa''. Struik Publishers Ltd, Cape Town (revised edition). * (1986): Phylogeny and classification of the Chamaeleonidae (Sauria) with special reference to hemipenis morphology. ''Bonner Zoologische Monographien'' 22: 1–64. * (1997): Liste der rezenten Amphibien und Reptilien - Chamaeleonidae. ''Das Tierreich'' 112: i-xiv, 1-85. * (2004): Phylogenetics of the southern African dwarf chameleons, ''Bradypodion'' (Squamata: Chamaeleonidae). ''Molecular Phylogenetics and Evolution'' 30: 354–365. PDF fulltext
{{Taxonbar, from=Q2323897 Lizard genera Taxa named by Leopold Fitzinger