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Anolis Carolinensis

TDactyloidae, Iguania, Sauria, Squamata or lizards 
S Anoles
LangE: North American Green Anole, Green anole
S: Anolis Verde
G: Rotkehlanolis 
NL: Roodkeelanolis
SynonymsAnolius carolinensis VOIGT in CUVIER & VOIGT 1832: 71
Lacerta principalis LINNAEUS 1758 (fide DUMÉRIL & BIBRON 1837: 121)
Anolis bullaris DAUDIN 1802: 69 (part.)
Agama bullaris — LINK 1807: 58
Agama strumosa — LINK 1807: 59
Anolis strumosa — HARLAN 1835: 143
Anolis Carolinensis — DUMÉRIL & BIBRON 1837: 120
Anolis podargicus RICHARDSON 1837: 200 (part.)
Dactyloa (Ctenocercus) carolinensis — FITZINGER 1843: 68
Anolis principalis — GRAY 1845: 202
Anolis baccatus BOCOURT 1873: 59
Anolis baccatus — BOULENGER 1885: 54
Anolis carolinensis — BOULENGER 1885: 43
Anolis principalis — LOENNBERG 1894
Anolis baccatus — SMITH & TAYLOR 1950
Anolis baccatus — FITCH & HENDERSON 1973: 127
Anolis carolinensis seminolus VANCE 1991: 75
Norops baccatus — LINER 1994
Anolis carolinensis — LINER 1994
Anolis carolinensis — MCKEOWN 1996
Norops baccatus — NICHOLSON 2002
Anolis carolinensis — NICHOLSON et al. 2005
Anolis baccatus — LINER 2007
Anolis carolinensis seminolus — COLLINS & TAGGART 2009
Anolis carolinensis seminolis [sic] — NICHOLSON et al. 2012
Anolis carolinensis seminolus — CROTHER et al. 2012
Anolis carolinensis — NICHOLSON et al. 2012
Anolis carolinensis — NICHOLSON et al. 2018
Anolis carolinensis seminolis — NICHOLSON et al. 2018 (in error) 
DistributionUSA (E Texas, SE Oklahoma, S Arkansas, Louisiana, Mississippi, Alabama, Georgia, Florida, South Carolina, North Carolina, SE Tennessee), Mexico (incl. Tamaulipas),
Bahamas, Grand Cayman Islands (HR 33: 321), Anguilla (HR 32: 118)

Introduced to Hawaii (fide MCKEOWN 1996) and California.
Introduced to Japan (Chichizima Is. and Hahazima Is. of Ogasawara Islands.And Okinawazima Is.).
Introduced to Micronesia and Guam (G. Rodda, pers. comm., 13 March 2016).
May have been introduced to Tenerife, Canary Islands, Spain (M. Lopez-Darias, Anole Annals July 20, 2016, Stroud et al. 2016)

baccatus: Mexico; Type locality: Mexico;

seminolus: USA (Florida); Type locality: 6.8 miles WNW of Murdock, Sarasota County, Florida.  
Reproductionoviparous Hybridization: Anolis porcatus appears to hybridize with A. carolinensis in Florida (T. Hagey, Anole Annals 2016). 
TypesNeotype: NCSM 93545 (North Carolina Museum of Natural Sciences); was ChR 862 (Charleston Museum), designated by Vance 1991, but collection was later moved to NCSM.
Holotype: MVZ (originally as UCMZ) 53793 [seminolus]
Holotype: MNHN 1126 [baccatus]
Synypes: MCZ 5955, 5956 [principalis] 
DiagnosisDiagnosis (Anolis s.s.): Support for this genus is provided by 47 apomorphies including seven morphological features and 40 molecular ones. There are two unequivocal morphological features: mental scale completely divided (26: a to z); and supratemporal processes leave supraocciptal exposed above (61: z to a). There are 23 unequivocal molecular apomorphies (see Appendix II, from NICHOLSON et al. 2012: 33).
 
CommentSynonymy: partly after VANCE 1991. BOULENGER 1885 listed A. porcatus as synonym of A. carolinensis. A specimen of A. baccatus reported from Sepaquite, Alta Verapaz, Guatemala by Barbour 1934: 124 is actually A. sericeus (fide Stuart, cited in SMITH & TAYLOR 1950: 62). ANDERSSON states that the Lacerta principalis of LINNAEUS 1758 is not Anolis carolinensis. Anolis cooperi BAIRD 1858 is a nomen dubium fide Vance, cited in Smith 1976: 252 (Synopsis VII).

Subspecies: VANCE 1991 described Anolis carolinensis seminolus, but more recent (genetic) studies didn’t find a clear overlap between mrophological seminolus and genetically defined populations, hence Crother et al. 2017 and others do not recognize subspecies within A. carolinensis.

Genetics: In July 2005, the scientific community overwhelmingly chose the green anole lizard, Anolis carolinensis, as its first target species for reptilian genome sequencing, with the American alligator, garter snake and/or painted turtle to follow (http://www.reptilegenome.org) (MODI & CREWS 2005). The genome sequence has been completed in 2011 (Alföldi et al. 2011). Partial genetic content of the Z chromosome is known in colubrids and viperids, and is highly syntenic to chromosome 6 (ACA6) of Anlis carolinensis (”ACA”, Rovatsos et al. 2015).

Evolution: On small islands in Florida, we found that the lizard Anolis carolinensis moved to higher perches following invasion by Anolis sagrei and, in response, adaptively evolved larger toepads after only 20 generations (Stuart et al. 2014).

Sexual dimorphism: the carolinensis and hendersoni clades are the most extreme both in male facial elongation and the degree of sexual dimorphism (Sanger et al. 2013).

Distribution: See maps in Vance 1991, Palmer & Braswell 1995: 116 (USA: Map 23). Probably erroneously reported from Belize. A single specimen (UF 23924) has been collected from Half Moon Cay (Belize), and Lee 1996 reports that this specimen indeed looks like A. carolinensis. Lee (2000) reports that efforts to find additional carolinensis on Half Moon Cay (Belize) have not been successful. However, Anolis allisoni is known from Belize (especially Half Moon Cay), and is very similar (and very closely related) to carolinensis. Not on Cuba (L. Mahler, pers. comm.). There are questionable records from SE Virginia (VANCE 1991).
In Japan, the green anole Anolis carolinensis invaded the Ogasawara Islands in 1960’s and Okinawa Island in 1980’s. Phylogenetic analysis shows that the invader A. carolinensis originated in the western part of the Gulf Coast and inland areas of the United States. Interestingly, all of the invaded A. carolinensis in Ogasawara, Okinawa and Hawaii originated from the Gulf Coast and inland areas of the United States (Suzuki-Ohno et al. 2017).

Species group: Anolis carolinensis species group (fide NICHOLSON et al. 2012).

Type species: Anolis bullaris DAUDIN 1802: 69 is the type species of the genus Anolis DAUDIN 1802 (fide SMITH & TAYLOR 1950: 56). Note however, that PETERS et al. 1970: 43 give Anolis bullaris Latreille as type species). See also Sabrosky 1983 and Stimson & Underwood 1983 for a discussion of the type species for Anolis.

Phylogenetics (genus). For a comprehensive phylogenetic analysis of anoles see Poe et al. 2017, 2018 and Román-Palacios et al. 2018.

Karyotype: 2n=36, XY (males) or XX (females) (Giovannotti et al. 2016) 
EtymologyNamed after the Carolinas where the species was found. A. c. seminolus has been named after the Seminole Indians of Florida.

The name Anolis is from the French l’anole, which is derived from anoli (or anolis) or anaoli (or anoali); aboriginal West Indian words meaning ‘‘lizard’’ (see Nicholson et al., 2012, for more information on the origin of Anolis). Anolis is masculine (Stimson & Underwood 1983). 
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Zootaxa 3626 (2): 295–299 – get paper herePoe, Steven; Adrián Nieto-Montes de Oca, Omar Torres-Carvajal, Kevin de Queiroz, Julián A. Velasco, Brad Truett, Levi N. Gray, Mason J. Ryan, Gunther Köhler, Fernando Ayala-Varela, and Ian Latella 2018. Comparative Evolution of an Archetypal Adaptive Radiation: Innovation and Opportunity in Anolis Lizards. The American Naturalist – get paper herePoe, Steven; Adrián Nieto-montes de Oca, Omar Torres-carvajal, Kevin De Queiroz, Julián A. Velasco, Brad Truett, Levi N. Gray, Mason J. Ryan, Gunther Köhler, Fernando Ayala-varela, Ian Latella 2017. A Phylogenetic, Biogeographic, and Taxonomic study of all Extant Species of Anolis (Squamata; Iguanidae). Syst Biol 2017: syx029 – get paper hereRajeev, M. & Rajeev, M. 2012. Geographic distribution: Anolis carolinensis (green anole). Herpetological Review 43: 304Rauh, J. 2006. Der Rotkehlanolis (Anolis carolinensis). Natur und Tier Verlag (Münster), 64 pp. – get paper hereRichardson, J. 1837. Report on North American Zoology. 6th Ann. Rept. British Assoc. 6: 121-224 [1836]Román-Palacios, C., Tavera, J., & Castañeda, M. del R. 2018. When did anoles diverge? An analysis of multiple dating strategies. Molecular Phylogenetics and Evolution, 127, 655–668 – get paper hereRovatsos M, Vukić J, Lymberakis P, Kratochvıl L. 2015. Evolutionary stability of sex chromosomes in snakes. Proc. R. Soc. B 282: 20151992 – get paper hereRovatsos, Michail; Marie Altmanová, Martina Pokorná and Lukáš Kratochvíl 2014. CONSERVED SEX CHROMOSOMES ACROSS ADAPTIVELY RADIATED ANOLIS LIZARDS. Evolution, DOI: 10.1111/evo.12357 – get paper hereSabrosky, C.W. 1983. Comment on the Type Species of Anolis Daudin, 1802. Bull. zool. Nomenclature 40 (1): 15-16 – get paper hereSanger, T. J., Sherratt, E., McGlothlin, J. W., Brodie, E. D., Losos, J. B. and Abzhanov, A. 2013. CONVERGENT EVOLUTION OF SEXUAL DIMORPHISM IN SKULL SHAPE USING DISTINCT DEVELOPMENTAL STRATEGIES. Evolution, 67: 2180–2193 – get paper hereSanger, Thomas J. & Bonnie K. Kircher 2017. Model Clades Versus Model Species: Anolis Lizards as an Integrative Model of Anatomical Evolution Methods in Molecular Biology book series (MIMB, volume 1650) – get paper hereSanger, Thomas J.; Liam J. Revell, Jeremy J. Gibson-Brown and Jonathan B. Losos 2012. Repeated modification of early limb morphogenesis programmes underlies the convergence of relative limb length in Anolis lizards Proc. R. Soc. B (2012) 279, 739–748, doi:10.1098/rspb.2011.0840 – get paper hereSater, N.M. & Smith, C.E. 2018. Anolis carolinensis (Green Anole) Interspecific mating. Herpetological Review 49 (1): 114-115. – get paper hereSchneider, C.J. 2009. Exploiting genomic resources in studies of speciation and adaptive radiation of lizards in the genus Anolis. Integrative and Comparative Biology, 48 (4): 520–526Sirsi, S., R., M.A., Forstner, M.R.J. & Fioley, D.H., III. 2017. Geographic Distribution: Anolis carolinensis (Green Anole). Herpetological Review 48 (4): 810-811 – get paper hereSmith,H.M. & Taylor,E.H. 1950. An annotated checklist and key to the reptiles of Mexico exclusive of the snakes. Bull. US Natl. Mus. 199: 1-253 – get paper hereStevenson, Dirk J., Christopher L. Jenkins, Kevin M. Stohlgren, John B. Jensen, David L. Bechler, Ian Deery, Daniel Duff, Sean P. Graham, Robb Herrington, Phillip Higgins, Robert V. Horan, III, Crystal Kelehear, Dylan Kelly, Kelsie Kincaid, Lance D. 2015. Significant new records of amphibians and reptiles from Georgia, USA. Herpetological Review 46 (4): 597-601 – get paper hereStimson, A.F. & Underwood, G.L. 1983. Comment on the Type Species of Anolis Daudin, 1802. Bull. zool. Nomenclature 40 (1): 17-19 – get paper hereStroud, Outerbridge, Giery 2016. First Specimen of an American Green Anole (Anolis carolinensis) on the Oceanic Island of Bermuda, with a Review of the Species’ Current Global Distribution. IRCF Rept Amph 23 (3):188–190 – get paper hereStuart, Y. E.; T. S. Campbell, P. A. Hohenlohe, R. G. Reynolds, L. J. Revell, J. B. Losos 2014. Rapid evolution of a native species following invasion by a congener. Science 346, 463 (2014);<br />DOI: 10.1126/science.1257008 – get paper hereStuart, Yoel E. 2010. Ecological character displacement in Anolis carolinensis. Anolis Newsletter VI: 186-193 – get paper hereSuzuki-Ohno Y, Morita K, Nagata N, et al. 2017. Factors restricting the range expansion of the invasive green anole Anolis carolinensis on Okinawa Island, Japan Ecol Evol. 2017;00:1–10 – get paper hereTerán-Juárez, Sergio A., Elí García Padilla, Vicente Mata-Silva, Jerry D. Johnson and Larry David Wilson. 2016. The herpetofauna of Tamaulipas, Mexico: composition, distribution, and conservation status. Mesoamerican Herpetology 3 (1): 43–113 – get paper hereTerán-Juárez, Sergio A.; Elí García-Padilla, Francisco E. Leyto-Delgado, and Leccinum J. García- Morales 2015. New records and distributional range extensions for amphibians and reptiles from Tamaulipas, Mexico. Mesoamerican Herpetology 2 (2): 208-214 – get paper hereThawley, Christopher J. and Fern Graves. 2014. Geographic Distribution: Anolis carolinensis (green anole). Herpetological Review 45 (2): 282 – get paper hereToda, Mitsuhiko; Noriyuki Komatsu, Hiroo Takahashi, Naomi Nakagawa, and Naozumi Sukigara 2013. Fecundity in Captivity of the Green Anoles, Anolis carolinensis, Established on the Ogasawara Islands. Current Herpetology Aug 2013, Vol. 32, No. 2: 82-88.Tollis M, Ausubel G, Ghimire D, Boissinot S 2012. Multi-Locus Phylogeographic and Population Genetic Analysis of Anolis carolinensis: Historical Demography of a Genomic Model Species. PLoS One 7 (6): e38474. doi:10.1371/journal.pone.0038474 – get paper hereTollis M, Boissinot S. 2014. Genetic variation in the green anole lizard (Anolis carolinensis) reveals island refugia and a fragmented Florida during the quaternary. Genetica. 2014 Feb;142(1):59-72. doi: 10.1007/s10709-013-9754-1Tollis, Marc & Stéphane Boissinot 2013. Genetic variation in the green anole lizard (Anolis carolinensis) reveals island refugia and a fragmented Florida during the quaternary. Genetica (2014) 142:59–72; DOI 10.1007/s10709-013-9754-1Unger, S D & Keenan, D M; 2018. Efficacy of visible implant elastomer (VIE) tag retention in the Green Anole, Anolis carolinensis, by using ImageJ color analysis Herpetological Review 49 (4): 677-681 – get paper hereValle, Luisa Dalla and Lorenzo Alibardi 2010. Forty hard keratin-associated beta-proteins (beta-keratins) allow the formation of all types of scales, adhesive pads and claws in Anolis carolinensis. Anolis Newsletter VI: 28-32 – get paper hereVance T. 1991. MORPHOLOGICAL VARIATION AND SYSTEMATICS OF THE GREEN ANOLE ANOLIS-CAROLINENSIS (REPTILIA, IGUANIDAE). Bull. Maryland Herp. Soc. 27 (2): 43-89. – get paper herevon Geldern, Charles E. 1919. Mechanism in the production of the throat-fan in the Florida chameleon, Anolis carolinensis. Proc. Cal. Acad. Sci. 9 (12): 313-329 – get paper herevon Geldern, Charles E. 1921. Color changes and structure of the skin of Anolis carolinensis. Proc. Cal. Acad. Sci. 10 (10): 77-117 + 3 plates – get paper hereWade, J. K.;Echternacht, A. C.;McCracken, G. F. 1983. Genetic variation and similarity in Anolis carolinensis (Sauria: Iguanidae). Copeia 1983 (2): 523-529 – get paper hereWalguarnery, Justin W.; Rachel M. Goodman, and Arthur C. Echternacht 2012. Thermal Biology and Temperature Selection in Juvenile Lizards of Co-occurring Native and Introduced Anolis Species. Journal of Herpetology 46 (4): 620-624. – get paper hereWard, Chelsea K. and Roger D. Birkhead. 2015. Anolis carolinensis (green anole) diet. Herpetological Review 46 (2): 250-251 – get paper hereWeller, W.H. 1930. Records of Some Reptiles and Amphibians from Chimney Rock Camp, Chimney Rock N. C., and Vicinity. Proc. Jr. Soc. Nat. Hist. Cincinnati 1 (8-9): 51-54 [unnumbered pages]Wilson, B. & Holt, B.D. 2017. Geographic Distribution: Anolis carolinensis (Green Anole). Herpetological Review 48 (4): 810 – get paper hereYabuta, Shinji and Akiko Suzuki-Watanabe 2011. Function of Body Coloration in Green Anoles (Anolis carolinensis) at the Beginning of the Breeding Season: Advertisement Signaling and Thermoregulation. Current Herpetology Dec 2011, Vol. 30, No. 2: 155-158. – get paper hereYasumiba, Kiyomi; Ayumi Okada, Isamu Okochi, and Noriko Iwai 2016. Minimum Longevity and Growth of the Invasive Green Anole, Anolis carolinensis, in Chichi-jima of the Ogasawara Islands, Japan Current Herpetology Aug 2016, Vol. 35, No. 2: 101-105. – get paper hereYuan, M. L., M. H. Wake, and I. J. Wang. 2019. Phenotypic integration between claw and toepad traits promotes microhabitat specialization in the Anolis adaptive radiation. Evolution 73:231–244 – get paper here 
External links
IUCN Red List – Anolis carolinensis – Least Concern, LCNational Center for Biotechnology Information

http://museum.nhm.uga.edu/~GAWildlife/Reptiles/reptsp.html
http://www.math.ie.kanagawa-u.ac.jp
http://gto.ncsa.uiuc.edu/pingleto/herps/lizards.html
http://www.wildherps.com/families/Polychrotidae.html
http://www.kingsnake.com/anolecare
http://www.anoleannals.org/2012/06/11/the-gray-dewlapped-anoleanolis-carolinensis-seminolus/
http://www.anoleannals.org/2014/05/30/green-anole-color-morphs/
http://www.spiegel.de/wissenschaft/natur/evolution-echse-entwickelt-klebefuesse-in-nur-15-jahren-a-998309.html
http://www.anoleannals.org/2015/06/10/are-brown-anoles-in-florida-really-driving-green-anoles-to-extinction/
http://www.anoleannals.org/2016/07/13/jmih-2016-genetic-evidence-of-hybridization-between-the-native-green-anole-anolis-carolinensis-and-the-invasive-cuban-green-anole-a-porcatus/
http://www.anoleannals.org/2016/07/20/introduced-anolis-species-in-tenerife-canary-islands-spain/
http://www.anoleannals.org/2017/05/19/factors-restricting-range-expansion-for-the-invasive-green-anole-anolis-carolinensis-on-okinawa-island-japan/Google images
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Anolis lizzard Genome

Comparative studies of amniotes have been hindered by a dearth of reptilian molecular sequences. With the genomic assembly of the green anole, Anolis carolinensis available, non-avian reptilian genes can now be compared to mammalian, avian, and amphibian homologs. Furthermore, with more than 350 extant species in the genus Anolis, anoles are an unparalleled example of tetrapod genetic diversity and divergence. As an important ecological, genetic and now genomic reference, it is imperative to develop a standardized Anolis gene nomenclature alongside associated vocabularies and other useful metrics.

Results

Here we report the formation of the Anolis Gene Nomenclature Committee (AGNC) and propose a standardized evolutionary characterization code that will help researchers to define gene orthology and paralogy with tetrapod homologs, provide a system for naming novel genes in Anolis and other reptiles, furnish abbreviations to facilitate comparative studies among the Anolis species and related iguanid squamates, and classify the geographical origins of Anolis subpopulations.

Conclusions

This report has been generated in close consultation with members of the Anolis and genomic research communities, and using public database resources including NCBI and Ensembl. Updates will continue to be regularly posted to new research community websites such as lizardbase. We anticipate that this standardized gene nomenclature will facilitate the accessibility of reptilian sequences for comparative studies among tetrapods and will further serve as a template for other communities in their sequencing and annotation initiatives.

Background

As the rate of generating new sequence assemblies continues to accelerate, the final bottleneck that remains is annotation. While automated pipelines have been developed, it is still up to community initiatives to pool, evaluate, integrate, and disseminate the necessary resources required for functional and comparative annotations that support research needs. The presence of multiple tools and resources, and changing assemblies and annotations, presents “moving-target” challenges for those attempting to assign function, orthology, nomenclature and other common vocabulary to genetic loci. One challenge is that many assemblies are, or will be, periodically updated due to resequencing efforts that aim to fill in ever-present gaps, initiatives to provide a consensus reference sequence that takes into account the polymorphism present in a species, or a re-deployment of different assembly algorithms. The second challenge is that the generation of confidently assigned gene models on a fixed assembly generally correlates with the amount of effort that a community puts into annotating their genome of interest. A third challenge relates to the principle that orthologous (and by association, functional) assignments are interdependent on the quality and quantity of annotations from closely related genomes.