Overview

Brief Summary

Anolis is a genus of lizards belonging to the family Polychrotidae. Anoles, as they are commonly known, are distributed from southeastern North America to central South America, including the West Indies. Most anoles are brown, grey or green, with a few exceptions including Anolis gorgonae, which is blue. Anoles are vertebrates (have a backbone or spine), have four limbs and a long tail, and are characterized by having adhesive toe-pads, and in most species, brightly colored throat fans, called dewlaps. Species found in the Caribbean are the most studied, but the majority of species are found in Central and South America (~60%). There is one anole species native to the United States, Anolis carolinensis, also known as the North American Green Anole (1). Often studied as an example of evolutionary diversification and adaptive radiation, the genus Anolis is an important research subject in areas like physiology, behavior, and community ecology, among others. The scientific importance of this group is reflected in that the North American green anole lizard was selected to be the first non-avian reptile to have its genome sequenced (3).

  • 1. Alföldi, J. et, al. The genome of the green anole lizard and a comparative analysis with birds and mammals. Nature 477, 587–591 (29 September 2011) doi:10.1038/nature10390
  • 2. Losos, J.B and Schneider, C. Anolis lizards Current Biology Vol 19 No 8 R316
  • 3. Sweetlove, L. Lizard genome unveiled. Nature. Published online 31 August 2011 doi:10.1038/news.2011.5123.
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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage

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Specimen Records: 1250
Specimens with Sequences: 1257
Specimens with Barcodes: 881
Species: 150
Species With Barcodes: 126
Public Records: 73
Public Species: 20
Public BINs: 59
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Statistics of barcoding coverage: Anolis sp. nov 1

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 0
Specimens with Barcodes: 2
Species With Barcodes: 1
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Statistics of barcoding coverage: Anolis fuscoauratus/scapularis

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 0
Specimens with Barcodes: 2
Species With Barcodes: 1
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Wikipedia

Norops (clade)

Not to be confused with Norops (genus)

Norops is a clade of 169 species of polychrotid lizards within the genus Anolis. Evidence for the clade's recognition stems from the fact that all of its designated species possess sufficient differences in morphology, especially in the structure and design of tail vertebrae, to suggest a deep divergence from the rest of Anolis.[1] The exact position of Norops and other distinct species groups within Anolis, either as clades or as genera, has been a subject of ongoing debate among anole scientists, however the description of Norops and other clades as genera is rapidly losing support within the scientific community;[2] however, it is unclear as to whether this debate will ever be completely resolved.

Taxonomic Status[edit]

In Richard E. Etheridge's original 1959 publication he recognized Anolis as an extremely diverse genus, and also recognized ten other genera including Norops, however he ultimately concluded that only four of these genera were morphologically unusual enough to warrant separation from Anolis.[3][4] Thus Etheridge concluded that the rest of the genera were simply deeply divergent groups of Anolis that were still firmly nestled within the genus. However the most recognized part of Etheridge's publication was his proposal to split Anolis into alpha and beta series based on morphological differences in caudal vertebrae.[4] In a 1986 publication Craig Guyer and Jay Savage applied the genus name Norops to Etheridge's beta anoles and also divided the rest of Anolis into four other genera. This proposal has been mostly rejected by the anole community as Norops was one of only a few of these genera proven to be monophyletic. During the 1990s and early 2000s some biologists[who?] working in Central America also began assigning certain species in Etheridge’s beta section of Anolis into Norops. Though Norops is usually proposed as a separate genus from Anolis the name Norops was originally treated by Kristen. E. Nicholson (2002) as a name for a clade of lizards nestled within the genus Anolis, but distinct in terms of certain aspects of skeletal morphology which differentiate them from all other anoles.[5] Ironically Nicholson is also the first author of a 2012 publication proposing the split of Anolis into several genera.[6] The classification of Norops as a clade is often preferred by the scientific community as classification of Norops as a genus separate from Anolis would necessitate the designation of several other species formerly placed in Anolis into newly created genera (as proposed by Guyer and Savage (1986) and Nicholson (2012)), several of which have proven to be paraphyletic; however the separation of Anolis into several distinct but interrelated clades would make anole classification easier without necessitating the creation of several new genera. This was also part of the reason why Etheridge left his beta anoles within the genus Anolis even though he recognized that there were some differences between the two groups.[citation needed] There is still much debate regarding the placement of Norops and other distinct groups within Anolis. In December 2012 the online website The Reptile Database revised their classification of Anolis and followed Nicholson in splitting the genus into eight new genera; however all the new genera were grouped back into Anolis in April 2013. Norops continues to be recognized as a genus by workers in Central America [2] However, it is universally rejected by West Indian workers. These differences may be due[according to whom?] to the fact the mainland fauna is dominated by only two deeply divergent clades (Norops and Dactyloa in Guyer and Savage's terminology), whereas the West Indian radiation is much more taxonomically diverse, and relationships less certain.

Morphological characteristics[edit]

Whether Norops is classified as a clade or a genus, most taxonomists have agreed on what morphological characteristics warrant the placement of a species inside the group. Etheridge classified his beta anoles as species that shared the certain aspects of vertebral morphology; Nicholson et al. went further by defining Norops as species that shared the following specific aspects of skeletal morphology:[6]

  1. structure of caudal vertebrae and mechanism of autotomy
  2. location of pineal foramen
  3. absence of pterygoid teeth
  4. usual absence of splenial
  5. lower jaw morphology
  6. number of lumbar vertebrae

Nicholson also used these criteria when proposing species to be placed in the genus Norops. There is remarkable morphological variation within this clade, even relative to other clades in Anolis; however most Norops species share a set of similar characteristics that are readily observable even to the untrained eye; they are usually small to mid-size anoles, though a few species such as A.garmani can grow much larger.[6] They typically have robust bodies and stout, blunt heads; they exist in a variety of ecomorphs.[6] Females rarely possess a dewlap.

List of Species.[edit]

The following is a list of species that have been proposed as belonging in the clade Norops.[7]

Anolis rodriguez

Phenotypical Comparison[edit]

Below is a table showing a phenotypical comparison of selected species in the proposed clade Norops.

species nameimage
Cuban brown anole (Anolis sagrei)
Anolis sagrei sagrei (displaying).jpg
Neotropical green anole (Anolis biporcatus)
Neotropical green anole.jpg
Graham's anole (Anolis grahami)
Graham's anole on post.jpg
Golfo-Dulce anole (Anolis polylepis)
Anolis polylepis, Costa Rica.JPG
Garland anole (Anolis vittigerus)
Norops vittigerus01.jpg
Jamaican giant anole (Anolis garmani)
Jamaika-anolis-22.jpg
Humble anole (Anolis humilis)
Anolis humilis.jpg
Cayman blue throated anole (Anolis conspersus)
Anolis conspersus.jpg
Sagua de Tánamo anole (Anolis rubribarbus)
Sagua de Tánamo anole.jpg
Silky anole (Anolis sericeus)
Norops sericeus.jpeg
Stream anole (Anolis poecilopus)
Stream Anole Norops poecilopus.jpg
Blemished anole (Anolis mariarum)
Norops mariarum.jpg
Spanish flag anole (Anolis allogus)
Spanish flag anole.jpg

Distribution and colonization[edit]

Brown anole (Anolis sagrei)

Norops is an extremely successful clade of anoles; it is one of two anole clades found on the mainland of South and Central America, the other being Dactyloa. The clade is especially prevalent in Central America which has been suggested as the reason why Central American scientists usually recognize it as a genus. They are also found on West Indian islands, including all four Greater Antillean islands and several islands in the Lesser Antilles. They are exceptionally good colonizers and several species have established thriving populations outside of their native territories, often outcompeting indigenous lizards, including other anoles. A few such examples are the Cuban brown anole, which is native to the Greater Antillean island of Cuba as well as the Bahamas, but has colonized numerous other Caribbean islands as well as certain areas of the continental United States, and has now spread as far east as Taiwan.[8] In parts of Florida the introduced brown anoles are having a negative effect on the native green anole population.[9] Green anoles were originally generally found anywhere from the hedges to the treetops, however, since the introduction of Brown anoles in the 1970s green anoles have been relegated almost exclusively to the treetops, while brown anoles have entirely colonized ground level areas.[9]

Another example is the Jamaican anole, also called Graham's anole (Anolis grahami) which was originally endemic to Jamaica but was deliberately introduced to Bermuda in 1905 to control the fruit fly population. However, the anole quickly outcompeted Bermuda's only native lizard, the Bermudan rock skink, and became the most widespread reptile species on the island.[10] It has further endangered the already vulnerable rock skink and inhibited the colonization of Bermuda by other anole species.[10] It is also believed that mainland species of Norops are the descendants of West Indian colonizers.[citation needed]

Origin[edit]

Graham's anole (Anolis Grahami) in Jamaica

Until recently[when?] it was believed that colonization from the mainland to the West Indies was responsible for the current distribution of the Norops clade across the Caribbean. This hypothesis was chosen because successful colonization from small islands to the mainland rarely occurs while the opposite is often true; however a paper published by Nicholson and colleagues concluded that the mainland Norops species were actually the result of speciation events occurring within populations of West Indian species that had made it to the mainland. This conclusion was drawn fron the fact that the Norops clade is nested within a branch of Anolis that is primarily West Indian; also mainland Norops species form a monophyletic clade that is nested within a group whose other members are West Indian.[11] The paper also asserted that a West Indian origin for Norops is entirely feasible as such island-to-mainland colonization events have occurred multiple times within Anolis.[11] Like most aspects of anole research, the scientific community's position on the origin of Norops is subject to debate.

References[edit]

  1. ^ Kristen E.Nicholson (2002). "Phylogenetic analysis and a test of the current infrageneric classification of Norops (beta Anolis)". Herpetological Monographs (The Herpetologists' League) 16 (1): 93–120. doi:10.1655/0733-1347(2002)016[0093:PAAATO]2.0.CO;2. 
  2. ^ a b Jonathan Lossos (January 7, 2011). "Norops last stand". Anole Annals.org. WordPress. Retrieved 19 May 2013. 
  3. ^ Etheridge, R. (“1959”, 1960) The Relationships of the Anoles (Reptilia: Sauria: Iguanidae): An Interpretation Based on Skeletal Morphology. PhD. Dissertation, University of Michigan, Ann Arbor, 236 pp.
  4. ^ a b Rich Glor (October 1, 2012). "Historical Perspective On Anole Genera". Anole Annals.org. WordPress. Retrieved 19 May 2013. 
  5. ^ Kevin de Queiroz (October 4, 2012). "The PhyloCode and the Names of Anole Clades". Anole Annals. WordPress. Retrieved 31 May 2013. 
  6. ^ a b c d Nicholson, Kristen A.; Brian I.Crother; Craig Guyer; Jay M.Savage (2012). "It is time for a new classification of anoles (Squamata:Dactyloidae)". Zootaxa (Magnolia Press): 1–108. 
  7. ^ "Norops Search results". The Reptile Database. Retrieved 27 May 2013. 
  8. ^ "Anolis sagrei (brown anole) disrtibution". Smithsonian Marine Station. June 12, 2007. Retrieved 19 May 2013. 
  9. ^ a b "Nonnatives - Brown Anole". Florida Fish and Wildlife Conservation Commission. Retrieved 19 May 2013. 
  10. ^ a b Cecilia Hackett (1986). "Lizards Of Bermuda". Bermuda Zoological Society. p. 5. Retrieved 20 May 2013. 
  11. ^ a b Nicholson, Kirsten E.; Glor, Richard E.; Kolbe, Jason J.; Larson, Allan; Blair Hedges, S.; Losos, Jonathan B. (2005). "Mainland colonization by island lizards". Journal of Biogeography 32 (6): 929–938. doi:10.1111/j.1365-2699.2004.01222.x. 
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Anolis

Anolis, or anoles, is a genus of iguanian (anole) lizards belonging to the family Dactyloidae. With 387 species, Anolis represents the world's most species-rich amniote (terrestrial vertebrates) genus.[1]

Taxonomy[edit]

This very large genus displays considerable paraphyly, but phylogenetic analysis suggests a number of subgroups or clades.[1][2]

These include;

Several species of Anolis are occasionally ascribed to the genus Norops, but the validity of the Norops genus is not widely accepted. The green anole (Anolis carolinensis) has recently become the first reptile to have its complete genome sequenced.[3]

Ecomorphs[edit]

Main article: Anolis ecomorph

Anolis lizards are some of the best examples of both adaptive radiation and convergent evolution. Populations of lizards on isolated islands diverge to occupy separate ecological niches, mostly in terms of the location within the vegetation where they forage (such as in the crown of trees vs. the trunk vs. underlying shrubs).[4] These divergences in habitat are accompanied by morphological changes primarily related to moving on the substrate diameter they most frequently encounter, with twig ecomorphs having short limbs, while trunk ecomorphs have long limbs.

In addition, these patterns repeat on numerous islands, with animals in similar habitats converging on similar body forms repeatedly.[5][4] This demonstrates adaptive radiation can actually be predictable based on habitat encountered, and experimental introductions onto formerly lizard-free islands have proven Anolis evolution can be predicted.[6][7][8][9]

Species[edit]

See also[edit]

References[edit]

  1. ^ a b Kristen A. Nicholson; Brian I. Crother; Craig Guyer; Jay M. Savage (11 September 2012). "It is time for a new clasification of anoles (Squamata:Dactyloidae)". Zootaxa. Magnolia Press. p. 38. Retrieved 19 May 2013. 
  2. ^ RICHARD E. GLOR, JONATHAN B. LOSOS and ALLAN LARSON. Out of Cuba: overwater dispersal and speciation among lizards in the Anolis carolinensis subgroup.Molecular Ecology (2005) 14, 2419–2432. doi: 10.1111/j.1365-294X.2005.02550.x
  3. ^ Anolis Genome Sequencing Project, Broad Institute
  4. ^ a b Losos, J.B. (2007). Detective work in the West Indies: integrating historical and experimental approaches to study island lizard evolution. BioScience 57:585-597.
  5. ^ Losos, J. B., Jackman, T. R., Larson, A., de Queiroz, K., & Rodriguez-Schettino, L. (1998) Contingency and determinism in replicated adaptive radiations of island lizards. Science, 279, 2115-2118.
  6. ^ Calsbeek, R. (2008). Experimental evidence that competition and habitat use shape the individual fitness surface. Journal of Evolutionary Biology 22:97-108.
  7. ^ Calsbeek, R., W. Buermann, and T.B. Smith. (2009). Parallel shifts in ecology and natural selection in an island lizard. BMC Evolutionary Biology 9:3.
  8. ^ Calsbeek, R., and R.M. Cox. (2010). Experimentally assessing the relative importance of predation and competition as agents of selection. Nature 465:613-616.
  9. ^ Calsbeek, R., and T.B. Smith. (2007). Probing the adaptive landscape using experimental islands: density-dependent natural selection on lizard body size. Evolution 61:1052-1061.

Further reading[edit]

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