Biology:Gecko

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Short description: Lizard belonging to the infraorder Gekkota

Gecko
Temporal range: 110–0 Ma
Albian – present
Phelsuma l. laticauda.jpg
Gold dust day gecko
Scientific classification e
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Squamata
Clade: Gekkonomorpha
Infraorder: Gekkota
Cuvier, 1817
Subgroups

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 1.6 to 60 centimetres (0.6 to 23.6 inches).

Geckos are unique among lizards for their vocalisations, which differ from species to species. Most geckos in the family Gekkonidae use chirping or clicking sounds in their social interactions. Tokay geckos (Gekko gecko) are known for their loud mating calls, and some other species are capable of making hissing noises when alarmed or threatened. They are the most species-rich group of lizards, with about 1,500 different species worldwide.[1]

All geckos, except species in the family Eublepharidae lack eyelids; instead, the outer surface of the eyeball has a transparent membrane, the brille. They have a fixed lens within each iris that enlarges in darkness to let in more light. Since they cannot blink, species without eyelids generally lick their own brilles when they need to clear them of dust and dirt, in order to keep them clean and moist.[2]

Unlike most lizards, geckos are usually nocturnal[3] and have excellent night vision; their colour vision in low light is 350 times more sensitive than human eyes.[4] The nocturnal geckos evolved from diurnal species, which had lost the rod cells from their eyes. The gecko eye, therefore, modified its cone cells that increased in size into different types, both single and double. Three different photo-pigments have been retained, and are sensitive to ultraviolet, blue, and green. They also use a multifocal optical system that allows them to generate a sharp image for at least two different depths.[5][6] While most gecko species are nocturnal, some species are diurnal and active during the day, which have evolved multiple times independently.[3]

Many species are well known for their specialised toe pads, which enable them to grab and climb onto smooth and vertical surfaces, and even cross indoor ceilings with ease. Geckos are well known to people who live in warm regions of the world, where several species make their home inside human habitations. These, for example the house gecko, become part of the indoor menagerie and are often welcomed, as they feed on insect pests; including moths and mosquitoes. Like most lizards, geckos can lose their tails in defence, a process called autotomy; the predator may attack the wriggling tail, allowing the gecko to escape.[7]

The largest species, Gigarcanum delcourti, is only known from a single, stuffed specimen probably collected in the 19th century found in the basement of the Natural History Museum of Marseille in Marseille, France. This gecko was 600 millimetres (24 inches) long, and it was likely endemic to New Caledonia, where it lived in native forests.[8] The smallest gecko, the Jaragua sphaero, is a mere 16 millimetres (0.63 inches) long, and was discovered in 2001 on a small island off the coast of Hispaniola.[9]

Etymology

The Neo-Latin gekko and English 'gecko' stem from Indonesian-Malaysian gēkoq,[10] it is a Malay word borrowed from Javanese,[11] from tokek, which imitates the sounds that some species like Tokay gecko make.[10][12]:120[13]:253

Common traits

Like other reptiles, geckos are ectothermic,[14] producing very little metabolic heat. Essentially, a gecko's body temperature is dependent on its environment. Also, to accomplish their main functions; such as locomotion, feeding, reproduction, etc., geckos must have a relatively elevated temperature.[14]

Shedding or molting

File:Leopard Gecko Shedding Skin.ogv All geckos shed their skin at fairly regular intervals, with species differing in timing and method. Leopard geckos shed at about two- to four-week intervals. The presence of moisture aids in the shedding. When shedding begins, the gecko speeds the process by detaching the loose skin from its body and eating it.[15] For young geckos, shedding occurs more frequently, once a week, but when they are fully grown, they shed once every one to two months.[16]

Adhesion ability

Close-up of the underside of a gecko's foot as it walks on vertical glass

About 60% of gecko species have adhesive toe pads which allow them to adhere to most surfaces without the use of liquids or surface tension. Such pads have been gained and lost repeatedly over the course of gecko evolution.[17] Adhesive toepads evolved independently in about eleven different gecko lineages, and were lost in at least nine lineages.[17]

It was previously thought that the spatula-shaped setae arranged in lamellae on gecko footpads enable attractive van der Waals' forces (the weakest of the weak chemical forces) between the β-keratin lamellae / setae / spatulae structures and the surface.[18][19] These van der Waals interactions involve no fluids; in theory, a boot made of synthetic setae would adhere as easily to the surface of the International Space Station as it would to a living-room wall, although adhesion varies with humidity.[20][21] However, a 2014 study suggests that gecko adhesion is in fact mainly determined by electrostatic interaction (caused by contact electrification), not van der Waals or capillary forces.[22]

The setae on the feet of geckos are also self-cleaning, and usually remove any clogging dirt within a few steps.[23][24][25] Polytetrafluoroethylene (PTFE), which has very low surface energy,[26] is more difficult for geckos to adhere to than many other surfaces.

Gecko adhesion is typically improved by higher humidity,[20][21][27][28][29] even on hydrophobic surfaces, yet is reduced under conditions of complete immersion in water. The role of water in that system is under discussion, yet recent experiments agree that the presence of molecular water layers (water molecules carry a very large dipole moment) on the setae, as well as on the surface, increase the surface energy of both, therefore the energy gain in getting these surfaces in contact is enlarged, which results in an increased gecko adhesion force.[20][21][27][28][29] Moreover, the elastic properties of the b-keratin change with water uptake.[20][21][27]

Gecko toes seem to be double-jointed, but this is a misnomer, and is properly called digital hyperextension.[30] Gecko toes can hyperextend in the opposite direction from human fingers and toes. This allows them to overcome the van der Waals force by peeling their toes off surfaces from the tips inward. In essence, by this peeling action, the gecko separates spatula by spatula from the surface, so for each spatula separation, only some force necessary. (The process is similar to removing Scotch Tape from a surface.)

Geckos' toes operate well below their full attractive capabilities most of the time, because the margin for error is great depending upon the surface roughness, and therefore the number of setae in contact with that surface.

Use of small van der Waals force requires very large surface areas; every square millimetre of a gecko's footpad contains about 14,000 hair-like setae. Each seta has a diameter of 5 μm. Human hair varies from 18 to 180 μm, so the cross-sectional area of a human hair is equivalent to 12 to 1300 setae. Each seta is in turn tipped with between 100 and 1,000 spatulae.[23] Each spatula is 0.2 μm long[23] (one five-millionth of a metre), or just below the wavelength of visible light.[31]

The setae of a typical mature 70-gram (2.5-ounce) gecko would be capable of supporting a weight of 133 kilograms (293 pounds):[32][33] each spatula could exert an adhesive force of 5 to 25 nN.[27][34] The exact value of the adhesion force of a spatula varies with the surface energy of the substrate to which it adheres. Recent studies[29][35] have moreover shown that the component of the surface energy derived from long-range forces, such as van der Waals forces, depends on the material's structure below the outermost atomic layers (up to 100 nm beneath the surface); taking that into account, the adhesive strength can be inferred.

Apart from the setae, phospholipids; fatty substances produced naturally in their bodies, also come into play.[36] These lipids lubricate the setae and allow the gecko to detach its foot before the next step.

The origin of gecko adhesion likely started as simple modifications to the epidermis on the underside of the toes. This was recently discovered in the genus Gonatodes from South America.[37][38] Simple elaborations of the epidermal spinules into setae have enabled Gonatodes humeralis to climb smooth surfaces and sleep on smooth leaves.

Biomimetic technologies designed to mimic gecko adhesion could produce reusable self-cleaning dry adhesives with many applications. Development effort is being put into these technologies, but manufacturing synthetic setae is not a trivial material design task.

Skin

Gecko skin does not generally bear scales, but appears at a macro scale as a papillose surface, which is made from hair-like protuberances developed across the entire body. These confer superhydrophobicity, and the unique design of the hair confers a profound antimicrobial action. These protuberances are very small, up to 4 microns in length, and tapering to a point.[39] Gecko skin has been observed to have an anti-bacterial property, killing gram-negative bacteria when they come in contact with the skin.[40]

The mossy leaf-tailed gecko of Madagascar, U. sikorae, has coloration developed as camouflage, most being greyish brown to black, or greenish brown, with various markings meant to resemble tree bark; down to the lichens and moss found on the bark. It also has flaps of skin, running the length of its body, head and limbs, known as the dermal flap, which it can lay against the tree during the day, scattering shadows, and making its outline practically invisible.[41]

Teeth

Geckos are polyphyodonts, and able to replace each of their 100 teeth every 3 to 4 months.[42] Next to the full grown tooth there is a small replacement tooth developing from the odontogenic stem cell in the dental lamina.[43] The formation of the teeth is pleurodont; they are fused (ankylosed) by their sides to the inner surface of the jaw bones. This formation is common in all species in the order Squamata.

Taxonomy and classification

Pores on the skin are often used in classification.

The infraorder Gekkota is divided into seven families, containing about 125 genera of geckos, including the snake-like (legless) pygopods.[17][44][45][46][47][3][48]

Legless lizards of the family Dibamidae, also referred to as blind lizards,[49] have occasionally been counted as gekkotans, but recent molecular phylogenies suggest otherwise.[50][51]

Gekkota
 
 

Diplodactylidae

 
 
 

Carphodactylidae

 
 

Pygopodidae

 
 
 
 
 

Eublepharidae

 
 
 

Sphaerodactylidae

 
 
 

Phyllodactylidae

 
 

Gekkonidae

 
 
 
 
 

Evolutionary history

Skeleton of Eichstaettisaurus, thought to be an early member of the gecko lineage
Fossil of Yantarogekko preserved in Baltic amber

Several species of lizard from the Late Jurassic have been considered early relatives of geckos, the most prominent and most well supported being the arboreal Eichstaettisaurus from the Late Jurassic of Germany. Norellius from the Early Cretaceous of Mongolia is also usually placed as a close relative of geckos.[52] The oldest known fossils of modern geckos are from the mid-Cretaceous Burmese amber of Myanmar (including Cretaceogekko), around 100 million years old, which have adhesive pads on the feet similar to those of living geckos.[53][54][55]

Species

Mediterranean house gecko

More than 1,850 species of geckos occur worldwide,[56] including these familiar species:

  • Coleonyx variegatus, the western banded gecko, is native to the southwestern United States and northwest Mexico.
  • Cyrtopodion brachykolon, the bent-toed gecko, is found in northwestern Pakistan ; it was first described in 2007.
  • Eublepharis macularius, the leopard gecko, is the most common gecko kept as a pet; it does not have adhesive toe pads and cannot climb the glass of a vivarium.
  • Gehyra mutilata (Pteropus mutilatus), the stump-toed gecko, is able to vary its color from very light to very dark to camouflage itself; this gecko is at home in the wild, as well as in residential areas.
  • Gekko gecko, the Tokay gecko, is a large, common, Southeast Asian gecko known for its aggressive temperament, loud mating calls, and bright markings.
  • Hemidactylus is genus of geckos with many varieties.
    • Hemidactylus frenatus, the common house gecko, thrives around people and human habitation structures in the tropics and subtropics worldwide.
    • Hemidactylus garnotii, the Indo-Pacific gecko, is found in houses throughout the tropics, and has become an invasive species of concern in Florida and Georgia in the US.
    • Hemidactylus mabouia, the tropical house gecko, Afro-American house gecko, or cosmopolitan house gecko, is a species of house gecko native to sub-Saharan Africa and also currently found in North, Central, and South America and the Caribbean.
    • Hemidactylus turcicus, the Mediterranean house gecko, is frequently found in and around buildings, and is an introduced species in the US.
  • Lepidodactylus lugubris, the mourning gecko, is originally an East Asian and Pacific species; it is equally at home in the wild and residential neighborhoods.
  • Pachydactylus bibroni, Bibron's gecko, is native to southern Africa; this hardy arboreal gecko is considered a household pest.
  • Phelsuma laticauda, the gold dust day gecko, is diurnal; it lives in northern Madagascar and on the Comoros. It is also an introduced species in Hawaii.
  • Ptychozoon is a genus of arboreal geckos from Southeast Asia also known as flying or parachute geckos; they have wing-like flaps from the neck to the upper leg to help them conceal themselves on trees and provide lift while jumping.
  • Rhacodactylus is genus of geckos native to New Caledonia.
    • Rhacodactylus ciliatus (now assigned to the genus Correlophus), the crested gecko, was believed extinct until rediscovered in 1994, and is gaining popularity as a pet.
    • Rhacodactylus leachianus, the New Caledonian giant gecko, was first described by Cuvier in 1829; it is the largest living species of gecko.
  • Sphaerodactylus ariasae, the dwarf gecko, is native to the Caribbean Islands; it is the world's smallest lizard.
  • Tarentola mauritanica, the crocodile or Moorish gecko, is commonly found in the Mediterranean region from the Iberian Peninsula and southern France to Greece and northern Africa; their most distinguishing characteristics are their pointed heads, spiked skin, and tails resembling those of a crocodile.

Reproduction

Most geckos lay a small clutch of eggs. Some are live-bearing and a few can reproduce asexually via parthenogenesis. Geckos also have a large diversity of sex-determining mechanisms including temperature-dependent sex determination and both XX/XY and ZZ/ZW sex chromosomes with multiple transitions among them over evolutionary time.[57] Madagascar day geckos engage in a mating ritual in which sexually mature males produce a waxy substance from pores on the back of their legs. Males approach females with a head swaying motion along with rapid tongue flicking in the female.[58]

Obligate parthenogenesis has evolved multiple times as a reproductive system in the family Gekkonidae.[59] It was shown in three different obligate parthenogenetic complexes of geckos that their oocytes are able to undergo meiosis. An extra premeiotic endoreplication of chromosomes is essential for obligate parthenogenesis in these geckos.[59] Appropriate segregation during meiosis to form viable progeny is facilitated by the formation of bivalents made from copies of identical chromosomes.

References

  1. ↑ "Search results – gecko". The Reptile Database. http://reptile-database.reptarium.cz/advanced_search?taxon=gecko&submit=Search. 
  2. ↑ Badger, David (2006). Lizards: a Natural History of Some Uncommon Creatures. St. Paul, MN: Voyageur Press. p. 47. ISBN 978-0760325797. 
  3. ↑ 3.0 3.1 3.2 Gamble, T.; Greenbaum, E.; Jackman, T.R.; Bauer, A.M. (August 2015). "Into the light: Diurnality has evolved multiple times in geckos". Biological Journal of the Linnean Society 115 (4): 896–910. doi:10.1111/bij.12536. 
  4. ↑ Roth, L.S.V.; Lundstrom, L.; Kelber, A.; Kroger, R.H.H.; Unsbo, P. (1 March 2009). "The pupils and optical systems of gecko eyes". Journal of Vision 9 (3): 27.1–11. doi:10.1167/9.3.27. PMID 19757966. 
  5. ↑ Roth, Lina S. V.; Lundström, Linda; Kelber, Almut; Kröger, Ronald H. H.; Unsbo, Peter (1 March 2009). "The pupils and optical systems of gecko eyes". Journal of Vision 9 (3): 27.1–11. doi:10.1167/9.3.27. PMID 19757966. 
  6. ↑ "Gecko-inspired multifocal contact lenses, cameras on the anvil". 8 May 2009. http://news.oneindia.in/2009/05/08/geckoinspired-multifocal-contact-lenses-cameras-on-theanv.html. 
  7. ↑ Mihai, Andrei (9 September 2009). "Gecko tail has a mind of its own". ZME Science. http://www.zmescience.com/medicine/gecko-tail-has-a-mind-of-its-own/. 
  8. ↑ Heinicke, Matthew P.; Nielsen, Stuart V.; Bauer, Aaron M.; Kelly, Ryan; Geneva, Anthony J.; Daza, Juan D.; Keating, Shannon E.; Gamble, Tony (2023-06-19). "Reappraising the evolutionary history of the largest known gecko, the presumably extinct Hoplodactylus delcourti, via high-throughput sequencing of archival DNA" (in en). Scientific Reports 13 (1): 9141. doi:10.1038/s41598-023-35210-8. ISSN 2045-2322. PMID 37336900. 
  9. ↑ Piper, Ross (2007). Extraordinary Animals: an Encyclopedia of Curious and Unusual Animals. Westport, Conn.: Greenwood Press. p. 143. ISBN 978-0313339226. https://archive.org/details/extraordinaryani0000pipe. 
  10. ↑ 10.0 10.1 gecko, n., Oxford English Dictionary, Second edition, 1989; online version September 2011. Accessed 29 October 2011. Earlier version first published in New English Dictionary, 1898.
  11. ↑ Wilkinson, Richard James (1932). "ge'kok". ge'kok. I. Mytilene, Greece: Salavopoulos & Kinderlis. p. 337. https://nla.gov.au/nla.obj-60272771/view?partId=nla.obj-435905740#page/n346/. Retrieved 2022-12-10. 
  12. ↑ Siti Zaleha Mat Diah; Rosli Hashim; Yong Hoi Sen; Daicus Belabut; Syuhadah Dzarawi N.; Lim Boo Liat (2010). "Preliminary Survey of Lizards of Pantai Melawi, Bachok, Kelantan, Malaysia" (in en). Malaysian Journal of Science 29 (special issue): 117–120. doi:10.22452/mjs.vol29nosp.13. ISSN 2600-8688. https://ejournal.um.edu.my/index.php/MJS/article/view/7338. Retrieved 11 December 2022. 
  13. ↑ Marcellini, Dale (February 1977). "Acoustic and Visual Display Behavior of Gekkonid Lizards". American Zoologist 17 (1): 251–260. doi:10.1093/icb/17.1.251. https://academic.oup.com/icb/article/17/1/251/172256. Retrieved 2023-01-07. 
  14. ↑ 14.0 14.1 Girons, Hubert (August 1980). "Thermoregulation in Reptiles with Special Reference to the Tuatara and Its Ecophysiology Tuatara". Victoria University of Wellington Library. http://nzetc.victoria.ac.nz/tm/scholarly/tei-Bio24Tuat02-t1-body-d2.html. 
  15. ↑ "GeckoCare - shedding". http://www.GeckoCare.net/shedding.php. 
  16. ↑ "Crested geckos shedding". Buddy Genius. 5 July 2020. https://buddygenius.com/crested-geckos-shed/. 
  17. ↑ 17.0 17.1 17.2 Gamble, Tony; Greenbaum, Eli; Jackman, Todd R.; Russell, Anthony P.; Bauer, Aaron M. (27 June 2012). "Repeated Origin and Loss of Adhesive Toepads in Geckos". PLOS ONE 7 (6): e39429. doi:10.1371/journal.pone.0039429. PMID 22761794. Bibcode: 2012PLoSO...739429G. 
  18. ↑ "Scientific image – gecko toe". NISE Network. http://www.nisenet.org/scientific-images/gecko_toe. 
  19. ↑ Santos, Daniel; Spenko, Matthew; Parness, Aaron; Sangbae, Kim; Cutkosky, Mark (2007). "Directional adhesion for climbing: Theoretical and practical considerations". Journal of Adhesion Science and Technology 21 (12–13): 1317–1341. doi:10.1163/156856107782328399. http://www.brill.nl/journal-adhesion-science-and-technology. Retrieved 2012-02-04. "Gecko "feet and toes are a hierarchical system of complex structures consisting of lamellae, setae, and spatulae. The distinguishing characteristics of the gecko adhesion system have been described [as] (1) anisotropic attachment, (2) high pulloff force to preload ratio, (3) low detachment force, (4) material independence, (5) self-cleaning, (6) antiself sticking and (7) nonsticky default state. ... The gecko's adhesive structures are made from ß-keratin (modulus of elasticity [about] 2 GPa). Such a stiff material is not inherently sticky; however, because of the gecko adhesive's hierarchical nature and extremely small distal features (spatulae are [about] 200 nm in size), the gecko's foot is able to intimately conform to the surface and generate significant attraction using van der Waals forces.". 
  20. ↑ 20.0 20.1 20.2 20.3 Puthoff, J.B.; Prowse, M.; Wilkinson, M.; Autumn, K. (2010). "Changes in materials properties explain the effects of humidity on gecko adhesion". Journal of Experimental Biology 213 (21): 3699–3704. doi:10.1242/jeb.047654. PMID 20952618. 
  21. ↑ 21.0 21.1 21.2 21.3 Prowse, M.S.; Wilkinson, Matt; Puthoff, Jonathan B.; Mayer, George; Autumn, Kellar (2011). "Effects of humidity on the mechanical properties of gecko setae". Acta Biomaterialia 7 (2): 733–738. doi:10.1016/j.actbio.2010.09.036. PMID 20920615. 
  22. ↑ Izadi, H.; Stewart, K.M.E.; Penlidis, A. (9 July 2014). "Role of contact electrification and electrostatic interactions in gecko adhesion". Journal of the Royal Society Interface 11 (98): 20140371. doi:10.1098/rsif.2014.0371. PMID 25008078. "We have demonstrated that it is the CE-driven electrostatic interactions which dictate the strength of gecko adhesion, and not the van der Waals or capillary forces which are conventionally considered as the main source of gecko adhesion.". 
  23. ↑ 23.0 23.1 23.2 Hansen, W.R.; Autumn, K. (2005). "Evidence for self-cleaning in gecko setae". Proceedings of the National Academy of Sciences 102 (2): 385–389. doi:10.1073/pnas.0408304102. PMID 15630086. Bibcode: 2005PNAS..102..385H. "Setae occur in uniform arrays on overlapping lamellar pads at a density of 14,400 per mm2". 
  24. ↑ "How geckos stick to walls". http://www.lclark.edu/~autumn/dept/geckostory.html. 
  25. ↑ Xu, Quan; Wan, Yiyang; Hu, Travis Shihao; Liu, Tony X.; Tao, Dashuai; Niewiarowski, Peter H.; Tian, Yu; Liu, Yue et al. (20 November 2015). "Robust self-cleaning and micromanipulation capabilities of gecko spatulae and their bio-mimics". Nature Communications 6: 8949. doi:10.1038/ncomms9949. PMID 26584513. Bibcode: 2015NatCo...6.8949X. 
  26. ↑ "Why do the gecko's feet not stick to a teflon surface?". http://www.JustAnswer.com/questions/bwl6-feet-gecko-lizard-not-stick. [unreliable source?]
  27. ↑ 27.0 27.1 27.2 27.3 Huber, G.; Mantz, H.; Spolenak, R.; Mecke, K.; Jacobs, K.; Gorb, S.N.; Arzt, E. (2005). "Evidence for capillarity contributions to gecko adhesion from single spatula nanomechanical measurements". Proceedings of the National Academy of Sciences 102 (45): 16293–6. doi:10.1073/pnas.0506328102. PMID 16260737. Bibcode: 2005PNAS..10216293H. 
  28. ↑ 28.0 28.1 Chen, B.; Gao, H. (2010). "An alternative explanation of the effect of humidity in gecko adhesion: stiffness reduction enhances adhesion on a rough surface". International Journal of Applied Mechanics 2 (1): 1–9. doi:10.1142/s1758825110000433. Bibcode: 2010IJAM....2....1C. 
  29. ↑ 29.0 29.1 29.2 Loskill, P.; Puthoff, J.; Wilkinson, M.; Mecke, K.; Jacobs, K.; Autumn, K. (September 2012). "Macroscale adhesion of gecko setae reflects nanoscale differences in subsurface composition". Journal of the Royal Society Interface 10 (78): 20120587. doi:10.1098/rsif.2012.0587. PMID 22993246. 
  30. ↑ Russell, A.P. (1975). "A contribution to the functional analysis of the foot of the Tokay, Gekko gecko (Reptilia: Gekkonidae)". Journal of Zoology 176 (4): 437–476. doi:10.1111/j.1469-7998.1975.tb03215.x. 
  31. ↑ Autumn, Kellar; Sitti, M.; Liang, Y.A.; Peattie, A.M.; Hansen, W.R.; Sponberg, S.; Kenny, T.W.; Fearing, R. et al. (2002). "Evidence for van der Waals adhesion in gecko setae". Proceedings of the National Academy of Sciences 99 (19): 12252–12256. doi:10.1073/pnas.192252799. PMID 12198184. Bibcode: 2002PNAS...9912252A. 
  32. ↑ "Geckos can hang upside down carrying 40kg". http://www.physics.org/facts/gecko-really.asp. 
  33. ↑ Autumn, Kellar (29 September 2003). "How do gecko lizards unstick themselves as they move across a surface?". Scientific American. http://www.scientificamerican.com/article.cfm?id=how-do-gecko-lizards-unst. Retrieved 23 March 2013. 
  34. ↑ Lee, Haeshin; Lee, Bruce P.; Messersmith, Phillip B. (2007). "A reversible wet / dry adhesive inspired by mussels and geckos". Nature 448 (7151): 338–341. doi:10.1038/nature05968. PMID 17637666. Bibcode: 2007Natur.448..338L. 
  35. ↑ Loskill, P.; Haehl, H.; Grandthyll, S.; Faidt, T.; Mueller, F.; Jacobs, K. (November 2012). "Is adhesion superficial? Silicon wafers as a model system to study van der Waals interactions". Advances in Colloid and Interface Science 179–182: 107–113. doi:10.1016/j.cis.2012.06.006. PMID 22795778. 
  36. ↑ Hsu, P.Y.; Ge, L.; Li, X.; Stark, A.Y.; Wesdemiotis, C.; Niewiarowski, P.H.; Dhinojwala, A. (24 August 2011). "Direct evidence of phospholipids in gecko footprints and spatula-substrate contact interface detected using surface-sensitive spectroscopy". Journal of the Royal Society Interface 9 (69): 657–664. doi:10.1098/rsif.2011.0370. PMID 21865250. 
  37. ↑ Higham, T.E.; Gamble, T.; Russell, A.P. (2017). "On the origin of frictional adhesion in geckos: small morphological changes lead to a major biomechanical transition in the genus Gonatodes". Biological Journal of the Linnean Society 120 (3): 503–517. doi:10.1111/bij.12897. 
  38. ↑ Russell, A.P.; Baskerville, J.; Gamble, T.; Higham, T. (November 2015). "The evolution of digit form in Gonatodes (Gekkota: Sphaerodactylidae) and its bearing on the transition from frictional to adhesive contact in gekkotans". Journal of Morphology 276 (11): 1311–1332. doi:10.1002/jmor.20420. PMID 26248497. 
  39. ↑ Green, DW; Lee, KK; Watson, JA; Kim, HY; Yoon, KS; Kim, EJ; Lee, JM; Watson, GS et al. (25 January 2017). "High quality bioreplication of intricate nanostructures from a fragile Gecko skin surface with bactericidal properties". Scientific Reports 7: 41023. doi:10.1038/srep41023. PMID 28120867. Bibcode: 2017NatSR...741023G. 
  40. ↑ Watson, Gregory S.; Green, David W.; Schwarzkopf, Lin; Li, Xin; Cribb, Bronwen W.; Myhra, Sverre; Watson, Jolanta A. (2015). "A gecko skin micro/Nano structure – A low adhesion, superhydrophobic, anti-wetting, self-cleaning, biocompatible, antibacterial surface". Acta Biomaterialia 21: 109–122. doi:10.1016/j.actbio.2015.03.007. PMID 25772496. 
  41. ↑ Pianka, Eric R. (2006). Lizards: Windows to the Evolution of Diversity. Berkeley, CA: University of California Press. pp. 247. ISBN 0-520-24847-3. https://Archive.org/details/lizardswindowsto00pian. 
  42. ↑ "Mechanism of Tooth Replacement in Leopard Geckos – Developmental Biology Interactive". http://www.DevBio.Biology.gatech.edu/?page_id=3229. 
  43. ↑ Gregory R. Handrigan; Kelvin J. Leung; Joy M. Richman (2010). "Identification of putative dental epithelial stem cells in a lizard with life-long tooth replacement". Development 137 (21): 3545–3549. doi:10.1242/dev.052415. PMID 20876646. 
  44. ↑ Han, D.; Zhou, K.; Bauer, A.M. (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 (3): 353–368. doi:10.1111/j.1095-8312.2004.00393.x. 
  45. ↑ Gamble, T.; Bauer, A.M.; Greenbaum, E.; Jackman, T.R. (July 2008). "Out of the blue: A novel, trans-Atlantic clade of geckos (Gekkota, Squamata)". Zoologica Scripta 37 (4): 355–366. doi:10.1111/j.1463-6409.2008.00330.x. https://epublications.marquette.edu/bio_fac/872. Retrieved 2023-08-18. 
  46. ↑ Gamble, Tony; Bauer, Aaron M.; Greenbaum, Eli; Jackman, Todd R. (21 August 2007). "Evidence for Gondwanan vicariance in an ancient clade of gecko lizards". Journal of Biogeography 35: 88–104. doi:10.1111/j.1365-2699.2007.01770.x. https://epublications.marquette.edu/cgi/viewcontent.cgi?article=1758&context=bio_fac. Retrieved 30 September 2020. 
  47. ↑ Gamble, T.; Bauer, A.M.; Colli, G.R.; Greenbaum, E.; Jackman, T.R.; Vitt, L.J.; Simons, A.M. (February 2011). "Coming to America: Multiple Origins of New World Geckos". Journal of Evolutionary Biology 24 (2): 231–244. doi:10.1111/j.1420-9101.2010.02184.x. PMID 21126276. 
  48. ↑ Gamble, Tony; Greenbaum, Eli; Jackman, Todd R.; Russell, Anthony P.; Bauer, Aaron M. (June 27, 2012). "Repeated Origin and Loss of Adhesive Toepads in Geckos". PLOS ONE 7 (6): e39429. doi:10.1371/journal.pone.0039429. PMID 22761794. Bibcode: 2012PLoSO...739429G. 
  49. ↑ Myers, P.; R. Espinosa (2008). "Infraorder GekkotaInfraorder Gekkota (blind lizards, geckos, and legless lizards)". The Animal Diversity Web (online). http://animaldiversity.ummz.umich.edu/site/accounts/classification/Gekkota.html. 
  50. ↑ Townsend, Ted M.; Larson, Allan; Louis, Edward; Macey, J. Robert (1 October 2004). "Molecular Phylogenetics of Squamata: The Position of Snakes, Amphisbaenians, and Dibamids, and the Root of the Squamate Tree". Systematic Biology 53 (5): 735–757. doi:10.1080/10635150490522340. PMID 15545252. 
  51. ↑ Vidal, Nicolas; Hedges, S. Blair (October 2005). "The phylogeny of squamate reptiles (lizards, snakes, and amphisbaenians) inferred from nine nuclear protein-coding genes". Comptes Rendus Biologies 328 (10–11): 1000–1008. doi:10.1016/j.crvi.2005.10.001. PMID 16286089. https://comptes-rendus.academie-sciences.fr/biologies/articles/10.1016/j.crvi.2005.10.001/. 
  52. ↑ Tałanda, Mateusz (September 2018). Benson, Roger. ed. "An exceptionally preserved Jurassic skink suggests lizard diversification preceded fragmentation of Pangaea" (in en). Palaeontology 61 (5): 659–677. doi:10.1111/pala.12358. https://onlinelibrary.wiley.com/doi/10.1111/pala.12358. Retrieved 2022-06-17. 
  53. ↑ Arnold, E.N.; Poinar, G. (2008). "A 100 million year old gecko with sophisticated adhesive toe pads, preserved in amber from Myanmar (abstract)". Zootaxa. http://www.mapress.com/zootaxa/2008/f/z01847p068f.pdf. Retrieved August 12, 2009. 
  54. ↑ Fontanarrosa, Gabriela; Daza, Juan D.; Abdala, Virginia (April 2018). "Cretaceous fossil gecko hand reveals a strikingly modern scansorial morphology: Qualitative and biometric analysis of an amber-preserved lizard hand". Cretaceous Research 84: 120–133. doi:10.1016/j.cretres.2017.11.003. ISSN 0195-6671. http://dx.doi.org/10.1016/j.cretres.2017.11.003. 
  55. ↑ Bauer, A M (2019-07-01). "Gecko Adhesion in Space and Time: A Phylogenetic Perspective on the Scansorial Success Story". Integrative and Comparative Biology 59 (1): 117–130. doi:10.1093/icb/icz020. ISSN 1540-7063. PMID 30938766. 
  56. ↑ "THE REPTILE DATABASE". http://www.reptile-database.org/. 
  57. ↑ Gamble, Tony; Coryell, J.; Ezaz, T.; Lynch, J.; Scantlebury, D.; Zarkower, D. (2015). "Restriction site-associated DNA sequencing (RAD-seq) reveals an extraordinary number of transitions among gecko sex-determining systems". Molecular Biology and Evolution 32 (5): 1296–1309. doi:10.1093/molbev/msv023. PMID 25657328. 
  58. ↑ Fry, C. and C. Roycroft 2009. "Phelsuma madagascariensis " (On-line), Animal Diversity Web. Accessed March 24, 2021
  59. ↑ 59.0 59.1 Dedukh D, Altmanová M, Klíma J, Kratochvíl L. Premeiotic endoreplication is essential for obligate parthenogenesis in geckos. Development. 2022 Apr 1;149(7):dev200345. doi: 10.1242/dev.200345. Epub 2022 Apr 19. PMID 35388415

Further reading

External links

Wikidata ☰ {{{from}}} entry

es:Geco ja:ヤモリ no:Gekkoer zh:壁虎



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