Chameleons after the Flood

There is a single family of these popular color-shifting lizards: the Chamaeleonidae. This is an Old World family, with native populations found in Africa, Madagascar, certain Indian Ocean Islands, the Middle East, India, Pakistan, and Sri Lanka. There are numerous recent and historical introductions, including southern Europe, Florida, and Hawaii.

While there are no known intergeneric hybrids, and only a handful of anecdotal interspecific hybrids, a distinctive and highly adapted morphology supports the idea that chameleons make up a monobaramin (a single lineage either part of or the whole of a created kind). Close relationships to agamids and possibly iguanids may make it difficult to ascertain a holobaramin (the entirety of a recognizable created kind) at this time. Specific design features within chameleons include a ballistic tongue, color-changing iridophores in the skin, and disconjugate eye movements. Many species demonstrate ornamentation with horns, crests, and casques, particularly seen in males.

There is a limited fossil record for chameleons, compared to many other lizard families, so relationships within the group primarily rely on genetics and morphology. Within the secular model (e.g. Tolley et al. 2013), chameleons first appeared after the Gondwanan break-up, though there have been competing arguments regarding the origin of chameleons: Africa or Madagascar. Chameleon diversification into modern genera is believed to have occurred within the Eocene, with subsequent speciation within genera occurring from the Oligocene to the Miocene accompanying global climate changes. One typical genus, Chamaeleo, likely first emerged in central or southern Africa, speciating extensively and dispersing further than any other genus, throughout Africa and into Eurasia (Main et al. 2022). Because the family emerged after Gondwana, vicariance played little role in early diversification of genera, though it may have had a role in certain speciation scenarios later (Raxworthy et al. 2002; Macey et al. 2008). Oceanic dispersal (rafting) became key to chameleon biogeographic expansion to Madagascar and the Indian Ocean islands.

Within the creation model, chameleons are a poster child for post-Flood diversification and dispersal. We can easily reinterpret the secular model using the climatic tentpoles within the Cenozoic for the post-Flood interval between the Flood itself (ending near the K/Pg boundary) and the Ice Age. Faced with the continental rearrangement of the post-Flood world, chameleons (whether the family diverged from a common ancestor with agamids or were a distinct baraminic kind) left the Ark in the Middle East, moved into Africa for initial generic radiation, entrenching in southern Africa with extensive speciation there, and dispersing through rafting to Madagascar (leading to prolific diversification there) and elsewhere.

The earliest fossil chameleons are found in Miocene strata. All fossil chameleons are placed within genera still living today. Notable examples include:

Chamaeleo intermedius is a fossil chameleon from Miocene Kenya (Hillenius 1978). Rather than preserving fossil bones, this amazing calcitic cast clearly details the head and part of the back of the animal. It was found at the same site that produced the fossil ape Kenyapithecus wickeri. While this chameleon was initially thought to have a transitional morphology between one Chamaeleo group and a group that later was named Trioceros, subsequent analysis suggests it should be placed within the Chamaeleo chamaeleon group (Klaver 1981).

Calumma benovskyi was found in Miocene strata from Rusinga Island, Kenya (Čerňanský et al. 2020). The genus is today only known from Madagascar, which clearly evidences that the genus originated on the African mainland before dispersing overwater (though this particular fossil species does not appear to be basal to the Malagasy species).

European Chamaeleo: Several species of Chamaeleo are known from Miocene deposits in Germany, the Czech Republic, Switzerland, and Greece, indicating that this post-Flood period was subtropical at the time (Čerňanský 2010; Čerňanský 2011; Georgalis et al. 2016). Tentatively identified chameleon fossil remains have been reported from several different Miocene sites in Anatolia, Turkey (Georgalis et al. 2022). Chamaeleo likely emerged in central/southern Africa, with Europe being an early dispersal. At some point, these Chamaeleo species disappeared from Europe, leaving it chameleon-free until recent times.

? Chamaeleonidae indet.: Fragmented fossils from a Miocene northern India hominoid site (Sankhyan and Čerňanský 2016) have been interpreted as a possible chamaeleonid. Within a creation model, it could suggest an early Chamaeleo lineage branching off from the Middle East into Asia, though it may not necessarily be related to recent Asian Chamaeleo.

The living Chamaeleo chamaeleon is known from subfossil Holocene remains in Spain (Čerňanský 2010). Chamaeleo is also known from Pleistocene Israel and Lebanon (Georgalis et al. 2016). Fossil fragments from a Pliocene South African deposit suggest the presence of at least two different groups of chameleons at the time, though they cannot be definitively assigned to genera (Dollion et al. 2015).

The biogeographic, genetic, and fossil evidence indicates that the immediate post-Flood dispersion of chameleons from the Ark was into Africa, followed by radiation into different genera in the mid-Paleogene. The oldest diverging lineages include the Malagasy Brookesia, the Seychelles Archaius, and the African Rieppeleon, which may indicate the Ark kind (if it is family-level) was represented by a smaller ‘leaf chameleon’ morphotype.

Rieppeleon are the short-tailed leaf chameleons residing along the eastern coast of Africa, from Ethiopia south to Mozambique. There are currently three described species. Fisseha et al. (2013) noted that there are fewer micro-endemics in the genus compared to Rhampholeon, possibly because they are low-altitude chameleons, which may work against isolation.

Rhampholeon are pygmy chameleons found in eastern Africa (Kenya, Democratic Republic of Congo, Tanzania, and Mozambique) along with a region along the west-central coast in Gabon and Nigeria. This disjunction is likely caused by an originally wide distribution split by climatic change contracting the pan-African forests, “due to the gradual northern movement of the African continent and the closure of the Tethys Ocean” (Matthee et al. 2004). There are currently 25 described species. All are inconspicuous, with several being distinct leaf mimics.

Chamaeleo is found extensively in southern Africa, north along the western and eastern coasts, and north into southern Europe and the Middle East, with a disjunct species further east in India, Pakistan, and Sri Lanka. There are currently 14 described species. Genetic analysis suggests that a few species diverged early (~Oligocene), with most others diverging in the Miocene, likely associated with climatic changes. Most are arboreal forest or savanna dwellers. One species, the Namaqua chameleon (C. namaquensis) is fully terrestrial, living in very arid regions of southwestern Africa. The Asian chameleon, C. zeylanicus, diverged from the Arabian Chamaeleo (prior to speciation of C. arabicus and C. calyptratus within Arabia) during the Miocene, possibly dispersing overland with later disjunction due to extinction of connecting populations (Main et al. 2022). Overwater dispersal between Arabia and India has also been suggested (Macey et al. 2008). Note, of course, the possible Miocene fossils from northern India, mentioned above, which may or may not be related.

Bradypodion are the Southern African dwarf chameleons. There are currently 20 described species. Most are inconspicuous, though males can display vivid breeding colors. Divergence between the two main clades in Bradypodion likely occurred in the Miocene, with extensive species radiation via rapid adaptation during the Pleistocene likely triggered by fluctuating formations of distinct vegetation micro-habitats (Tolley et al. 2006; Tolley et al. 2008; Da Silva and Tolley 2013; Da Silva and Tolley 2017; Giles and Arbuckle 2022).

Kinyongia are the East African dwarf chameleons, found in Kenya, Tanzania, Uganda, and Rwanda. There are currently 23 described species. Genetic analysis suggests that diversification within the genus took place prior to the cooling that fragmented the pan-African forests (starting in the late Eocene, with additional climatic changes and aridification through to the Miocene), isolating species on separate montane forests (a form of vicariance) (Tolley et al. 2011).

Nadzikambia are the mountain dwarf chameleons of Malawi and Mozambique. There are currently two described species from sub-montane forests (Tilbury et al. 2006; Branch and Tolley 2010).

Trioceros are the equatorial chameleons of west-central and east-central equatorial Africa. There are currently 41 described species. Unlike other genera, they have managed to broadly maintain distribution across central Africa, suggesting the genus is more ecologically adaptable. Speciation may be due to localized vicariant events produced by geomorphological and climatic changes (Ceccarelli et al. 2014).

There are four genera of chameleons in Madagascar, with evidence supporting two primary overwater (rafting) dispersions from the African mainland. A third rafting dispersal from Africa led to the colonization of the Seychelles. Success in this form of dispersal appears to be supported by specific traits of the dispersing species: coastal distribution, larger body size (or at least large clutch size), and lineages with extreme (fastest or slowest) life histories (Weil et al. 2022).

The subfamily Brookesinae includes two genera, Brookesia and Palleon. These small lizards are known as leaf chameleons due to their morphological mimicry. There are currently 32 described species of Brookesia and two described species of Palleon. The Brookesinae are the earliest diverging lineage within the chameleon family (Tolley et al. 2013), and genetic analysis suggests that Palleon diverged very early (possibly Paleocene) from Brookesia (Glaw et al. 2013). This indicates that the Brookesinae lineage was an early breakaway lineage from the post-Ark African population that rafted to Madagascar. Parsimoniously, the two genera then diverged on Madagascar. Extensive micro-endemic speciation with Brookesia may be associated with Eocene-Oligocene climate shifts and forest fragmentation (Townsend et al. 2009).

Calumma includes the largest chameleon, by weight and length, the Parson’s chameleon, Calumma parsonii. There are currently 41 described species in the genus. Due to the Miocene fossil Calumma found on Rusinga Island, Kenya, we can ascertain that Calumma diverged on the African mainland, and dispersed separately to Madagascar from the Brookesinae (Čerňanský et al. 2020), then radiating extensively within Madagascar.

There are currently 24 described species of Furcifer. It shares a most recent common ancestor with Calumma (Tolley et al. 2013), indicating that it diverged from that genus once Calumma reached Madagascar. Speciation appears to have been driven by multiple drivers, including geographic barriers and niche partitioning, from the Miocene to the Pleistocene (Florio and Raxworthy 2016). Furcifer itself colonized two separate islands of the Comoros (found to the northwest of Madagascar) independently, leading to the emergence of a distinct species on each (Rocha et al. 2005). Interestingly, the genetic divergence of Furcifer cephalolepis on Grand Comoro is much higher than expected within the secular model, given how ‘recent’ (0.5 My) they believe that volcanic island to be.

The Seychelles tiger chameleon, Archaius tigris, is the sole species in its genus. Genetic analysis determined that it is a sister-taxon to Africa’s Rieppeleon, indicating an early overwater dispersal from the coast of Africa during the Palaeogene (Townsend et al. 2011).

Because of their unusual, yet attractive appearance, chameleons are popular pets and have been deliberately introduced into numerous ecosystems (Glaw 2015). Notable cases include:

Veiled chameleons (Chamaeleo calyptratus) were introduced to the Canary Islands (Pino-Vera et al. 2023)
Veiled chameleons were introduced to Hawaii, but eradicated in 2008 (Basso et al. 2019)
Veiled chameleons were introduced in Florida, multiple times at different locations (Krysko et al. 2004; Edwards et al. 2014)
Jackson’s chameleons (Trioceros jacksonii) were introduced to Hawaii (Chiaverano et al. 2014)
Panther chameleons (Furcifer pardalis) were introduced multiple times in southern Florida (Rochford et al. 2013; Fieldsend et al. 2021)
Panther chameleons were introduced to the islands of Mauritius and Reunion (Glaw 2015)
Oustalet’s chameleons (Furcifer oustaleti) were introduced to southern Florida (Smith et al. 2016)

Genetic evidence suggests that many (most? all?) modern-day European populations of the Mediterranean chameleon, Chamaeleo chamaeleon, were introduced in historical periods rather than being Plio-Pleistocene colonizations. Paulo et al. (2002) determined that two separate populations on the Iberian Peninsula were likely brought from different parts of North Africa at different times. Schembri (1983) noted that the chameleons on Malta were anecdotally introduced by Jesuit priests to a garden in St. Julians in the late 1800s, with genetic analysis pointing to two different origins for Maltese chameleons: Israel and Morocco (Andreone et al. 2016). Populations in Cyprus, Crete, and other islands have been traced genetically to Turkey, and those in mainland Italy are recently introductions from Tunisia and Lebanon (Andreone et al. 2016; Basso et al. 2019).

A second chameleon, the African chameleon, Chamaeleo africanus, has also been introduced into Greece. Specimens collected on the Greek mainland have haplotypes pointing to an origin in the Nile Delta (Kosuch et al. 1999; Dimaki et al. 2008).

Andreone, F., et al. 2016. The common chameleon Chamaeleo chamaeleon in southern Italy: Evidence for allochthony of populations in Apulia and Calabria (Reptilia: Squamata: Chamaeleonidae). Italian Journal of Zoology 83(3): 372-381.

Basso, R., et al. 2019. Multiple origins of the common chameleon in southern Italy. Herpetozoa 32: 11-19.

Branch, W. R., and K. A. Tolley. 2010. A new species of chameleon (Sauria: Chamaeleonidae: Nadzikambia) from Mount Mabu, central Mozambique. African Journal of Herpetology 59(2): 157-172.

Ceccarelli, F. S., et al. 2014. Evolutionary relationships, species delimitation and biogeography of Eastern Afromontane horned chameleons (Chamaeleonidae: Trioceros). Molecular Phylogenetics and Evolution 80: 125-136.

Čerňanský, A. 2010. A revision of chamaeleonids from the Lower Miocene of the Czech Republic with description of a new species of Chamaeleo (Squamata, Chamaeleonidae). Geobios 43: 605-613.

Čerňanský, A. 2011. A revision of the chameleon species Chamaeleo pfeili Schleich (Squamata; Chamaeleonidae) with description of a new material of chamaeleonids from the Miocene deposits of southern Germany. Bulletin of Geosciences 86(2): 275-282.

Čerňanský, A., et al. 2020. The only complete articulated early Miocene chameleon skull (Rusinga Island, Kenya) suggests an African origin for Madagascar’s endemic chameleons. Scientific Reports 10(109).

Chiaverano, L. M., M. J. Wright, and B. S. Holland. 2014. Movement behavior is habitat dependent in invasive Jackson’s chameleons in Hawaii. Journal of Herpetology 48(4): 471-479.

Da Silva, J. M., and K. A. Tolley. 2013. Ecomorphological variation and sexual dimorphism in a recent radiation of dwarf chameleons (Bradypodion). Biological Journal of the Linnean Society 109: 113-130.

Da Silva, J. M., and K. A. Tolley. 2017. Diversification through ecological opportunity in dwarf chameleons. Journal of Biogeography 44(4): 834-847.

Dimaki, M., A. K. Hundsdörfer, and U. Fritz. 2008. Eastern Mediterranean chameleons (Chamaeleo chamaeleon, Ch. africanus) are distinct. Amphibia-Reptilia 29: 535-540.

Dollion, A. Y., et al. 2015. Morphometric analysis of chameleon fossil fragments from the early Pliocene of South Africa: A new piece of the chamaeleonid history. Science of Nature 102(2): 1-14.

Edwards, J. R., et al. 2014. New county record for the veiled chameleon (Chamaeleo calyptratus Duméril and Bibron 1851), in Broward County, Florida, with notes on intentional introductions of chameleons in Southern Florida. Reptiles & Amphibians 21(2): 83-85.

Fieldsend, T. W., et al. 2021. Extreme male color polymorphism supports the introduction of multiple native-range panther chameleon (Furcifer pardalis) lineages to Florida, USA. Reptiles & Amphibians 28(2): 257-261.

Fisseha, M., J. Mariaux, and M. Menegon. 2013. The “Rhampoleon uluguruensis complex” (Squamata: Chamaeleonidae) and the taxonomic status of the pygmy chameleons in Tanzania. Zootaxa 3746(3): 439-453.

Florio, A. M., and C. J. Raxworthy. 2016. A phylogeographic assessment of the Malagasy giant chameleons (Furcifer verrucosus and Furcifer oustaleti). PLoS ONE 11(6): e0154144.

Georgalis, G. L., A. Villa, and M. Delfino. 2016. First description of a fossil chamaeleonid from Greece and its relevance for the European biogeographic history of the group. Science of Nature 103(12): 1-12.

Georgalis, G. L., et al. 2022. The antiquity of Asian chameleons—First potential Chamaeleonidae and associated squamate fauna from the lower and middle Miocene of Anatolia. Journal of Vertebrate Paleontology 42(2): e2160644.

Giles, S. A. W., and K. Arbuckle. 2022. Diversification dynamics of chameleons (Chamaeleonidae). Journal of Zoology 318: 241-252.

Glaw, F. 2015. Taxonomic checklist of chameleons (Squamata: Chamaeleonidae). Vertebrate Zoology 65(2): 167-246.

Glaw, F., O. Hawlitschek, and B. Ruthensteiner. 2013. A new genus name for an ancient Malagasy chameleon clade and a PDF-embedded 3D model of its skeleton. Salamandra 49(4): 237-238.

Hillenius, D. 1978. Notes on chameleons IV. A new chameleon, from the Miocene of Fort Ternan, Kenya (Chamaeleonidae, Reptilia). Beaufortia 343(28): 9-15.

Klaver, C. J. J. 1981. Lung-morphology in the Chamaeleonidae (Sauria) and its bearing upon phylogeny, systematics and zoogeography. Journal of Zoological Systematics and Evolutionary Research 19: 36-58.

Kosuch, J., M. Vences, and W. Böhme. 1999. Mitochondrial DNA sequence data support the allocation of Greek mainland chameleons to Chamaeleo africanus. Amphibia-Reptilia 20: 440-443.

Krysko, K. L., K. M. Enge, and F. W. King. 2004. The veiled chameleon, Chamaeleo calyptratus: A new exotic lizard species in Florida. Florida Scientist 67(4): 249-253.

Macey, J. R., et al. 2008. Socotra Island the forgotten fragment of Gondwana: Unmasking chameleon lizard history with complete mitochondrial genomic data. Molecular Phylogenetics and Evolution 49: 1015-1018.

Main, D. C., et al. 2022. Out of southern Africa: Origins and cryptic speciation in Chamaeleo, the most widespread chameleon genus. Molecular Phylogenetics and Evolution 175: 107578.

Matthee, C. A., C. R. Tilbury, and T. Townsend. 2004. A phylogenetic review of the African leaf chameleons: Genus Rhampholeon (Chamaeleonidae): The role of vicariance and climate change in speciation. Proceedings of the Royal Society of London B 271: 1967-1975.

Paulo, O. S., et al. 2002. The double origin of Iberian peninsular chameleons. Biological Journal of the Linnean Society 75: 1-7.

Pino-Vera, R., N. Abreu-Acosta, and P. Foronda. 2023. Study of zoonotic pathogens in alien population of veiled chameleons (Chamaeleo calyptratus) in the Canary Islands (Spain). Animals 13: 2288.

Raxworthy, C. J., M. R. J. Forstner, and R. A. Nussbaum. 2002. Chameleon radiation by oceanic dispersal. Nature 415: 784-787.

Rocha, S., M. A. Carretero, and D. J. Harris. 2005. Mitochondrial DNA sequence data suggests two independent colonizations of the Comoros archipelago by chameleons of the genus Furcifer. Belgian Journal of Zoology 135(1): 39-42.

Rochford, M. R., et al. 2013. The panther chameleon, Furcifer pardalis (Cuvier 1829) (Chamaeleonidae), another introduced chameleon species in Florida. Reptiles & Amphibians 20(4): 205-207.

Sankhyan, A. R., and A. Čerňanský. 2016. A first possible chameleon from the late Miocene of India (the hominoid site of Haritalyangar): A tentative evidence for an Asian dispersal of chameleons. The Science of Nature 103(94): 1-6.

Schembri, P. J. 1983. The Mediterranean chameleon. Civilization 9: 238-239.

Smith, D., et al. 2016. Observations on nesting and clutch size in Furcifer oustaleti (Oustalet’s chameleon) in South Florida. Southeastern Naturalist 15(Special Issue 8): 75-88.

Tilbury, C. R., K. A. Tolley, and W. R. Branch. 2006. A review of the systematics of the genus Bradypodion (Sauria: Chamaeleonidae), with the description of two new genera. Zootaxa 1363: 23-38.

Tolley, K. A., B. M. Chase, and F. Forest. 2008. Speciation and radiations track climate transitions since the Miocene Climatic Optimum: A case study of southern African chameleons. Journal of Biogeography 35: 1402-1414.

Tolley, K. A., T. M. Townsend, and M. Vences. 2013. Large-scale phylogeny of chameleons suggests African origins and Eocene diversification. Proceedings of the Royal Society B 280: 20130184.

Tolley, K. A., et al. 2006. Biogeographic patterns and phylogeography of dwarf chameleons (Bradypodion) in an African biodiversity hotspot. Molecular Ecology 15: 781-793.

Tolley, K. A., et al. 2011. Ancient forest fragmentation or recent radiation? Testing refugial speciation models in chameleons within an African biodiversity hotspot. Journal of Biogeography 38: 1748-1760.

Townsend, T. M., et al. 2009. Testing species-level diversification hypotheses in Madagascar: The case of microendemic Brookesia leaf chameleons. Systematic Biology 58(6): 641-656.

Townsend, T. M., et al. 2011. Eastward from Africa: Palaeocurrent-mediated chameleon dispersal to the Seychelles islands. Biology Letters 7: 225-228.

Weil, S.-S., et al. 2022. Chameleon biogeographic dispersal is associated with extreme life history strategies. Ecography e06323.

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zoocreation

Chameleons after the Flood

Chad Arment (2025)

Early Divergence and Dispersal

The African Chameleons

Bradypodion caeruleogula

Bradypodion pumilum

Chamaeleo calyptratus

Chamaeleo chameleon

Chamaeleo chameleon

Chamaeleo dilepis

Chamaeleo monachus

Chamaeleo namaquensis

Chamaeleo zeylanicus

Kinyongia msuyae

Kinyongia multituberculata

Kinyongia tavetana

Kinyongia tenuis

Nadzikambia baylissi

Nadzikambia mlanjensis

Rhampholeon spectrum

Rhampholeon tilburyi

Rieppeleon brachyurus

Rieppeleon brevicaudatus

Rieppeleon kerstenii

Trioceros cristatus

Trioceros deremensis

Trioceros jacksonii

Trioceros rudis

Madagascar and the Indian Ocean Islands

Archaius tigris

Brookesia vadoni

Brookesia perarmata

Brookesia superciliaris

Brookesia tuberculata

Calumma amber

Calumma crypticum

Calumma gallus

Calumma parsonii

Furcifer bifidus

Furcifer labordi

Furcifer pardalis

Furcifer campani

Furcifer lateralis

Furcifer minor

Furcifer willsii

Furcifer pardalis

Palleon nasus

Human Introduction

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2021-2025

zoocreation