Tupinambis

Tupinambis
Tupinambis teguixin
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Squamata
Family: Teiidae
Genus: Tupinambis
Daudin, 1802

Tupinambis is a lizard genus which belongs to the family Teiidae, and contains seven described species. These large lizards are commonly referred to as tegus (teiús in Portuguese); T. merianae (Argentine black and white tegu), T. rufescens (red tegu), and T. teguixin (gold tegu) are popular in the pet trade. They are primarily found in South America, although T. teguixin also occurs in Panama. Tegus that have escaped or have been illegally released have adapted and are increasing in several Florida counties including rural and suburban (especially south Miami-Dade and Hillsborough) counties, agricultural areas (especially Homestead and unincorporated Miami-Dade County, Florida) and publicly owned conservation areas (especially Southern Glades Environmental and Wildlife Area and southeast margin of Everglades National Park) of South Florida See invasive species website (www.IVE-GOT-1.org) for details with specific locations of credible observations and voucher specimens.

In 2012 a number of tegu species were reclassified from Tupinambis to the previously used genus Salvator. The newly proposed classification comes from a restructuring of the Teiidae family based upon the study of 137 morphological characteristics. The new classification is as follows: Salvator duseni (yellow tegu), Salvator rufescens (red tegu), Salvator merianae (black and white tegu), Tupinambis teguixin (gold tegu), Tupinambis longilineus (Rhondonia tegu), Tupinambis palustris (swamp tegu), Tupinambis quadrilineatus (four-lined tegu).[1]

Description

Lateral view of a red tegu skull, showing heterodont dentition.

Tegus are large reptiles, with some species reaching a total length of around 1.23 m (4.0 ft), and a weight of approximately 6.8 kg (15 lb). These opportunistic, wide-ranging lizards can be found in a variety of habitats, from swamps to rainforests to savannahs and cities.[2] Although terrestrial, they are capable swimmers, able to remain submerged for up to 22 minutes and having even been caught in gill nets set at sea.[3] Tupinambis have heterodont dentition consisting of four different types of teeth.[4] Incisor-type—tricuspid—teeth reside at the tip of the mouth.[4] Recurved canine-type teeth occur further back on the tooth row.[4] Behind those reside a separate set of incisor-like teeth (though flattened in a perpendicular plane to the first set of incisors).[4] The rearmost teeth are blunt, rounded, peg-shaped teeth.[4] The rearmost two tooth classes only occur in sexually mature individuals thus indicating an ontogenetic shift in tooth morphology.[4] Along with changes in tooth type, the frequency of each tooth type also changes with ontogeny, without an overall changes in tooth count (approximately 70 teeth).[4] Rather than increase tooth count, the teeth themselves increase in size as the jaw grows from hatchling to adult.[4] This ontogenetic shift in tooth morphology suggests a shift in diet with age, however few dietary studies have been done to support this claim and limited stomach content observations do not show much variability between hatchlings and juveniles.[4] Biomechanical studies have shown that Tegus have stronger limb bones than comparably-sized mammals or birds, a trait that may be inherent to amphibians and reptiles.[5] They exhibit social and maternal behaviour; female Tegus construct burrows to lay their eggs in, and will protect their brood until they hatch. Up to 35 eggs are produced in a clutch.[6] Tegus will hibernate together in groups, though males exhibit territorial behavior towards each other.[7] Tegus exhibit sexual dimorphism. Males display greatly expanded and prominent "jowls" along the base of the lower jaw. These jowls are the result of extensive growth of the ventral pterygoideus muscles (a prominent jaw closing muscle in reptiles).[8][9] Though both sexes exhibit hypertrophy of the ventral pterygoideus muscle, likely in response to their durophagous habits,[10] male Tupinambis show both greater relative and absolute muscle mass compared to females.[10] Further evidence for sexual selection of these jowls comes from observations that ventral pterygoideus muscle mass increases in males during the reproductive season.[11] Metabolic changes also occur during the reproductive season, in which the body temperature is increased up to 10 degrees and sustained internally like a bird or a mammal. This discovery has major evolutionary implications, providing support for the hypothesis that endothermy may have evolved in response to parental care. [12]

Ecology

Tegus are omnivorous, foraging for a wide range of foods using their forked tongues, including fruit, fungi, various arthropods, small vertebrates, carrion, and eggs.[13] The amount of meat that is consumed by tegus decreases as the animals mature.[14] As adults, tegus have few predators. Among them are big cats, birds of prey and large snakes.[15] Tegus defend themselves using their powerful jaws, which can exert forces of up to 1000N.[16] A bite from an adult tegu can crush human fingers.[17]

Though more terrestrial (morphologically less well-adapted for climbing into tree canopies or for swimming) tegus fill a similar ecological niche in South America to that filled by monitor lizards in Africa, Asia and Australia, and are an example of convergent evolution. Though similar in appearance to monitors, tegus are not closely related and can be distinguished by their larger heads, shorter necks, heavier bodies and different arrangement of the scales on the body and tail. Monitors have laterally compressed tails, well-suited for aquatic propulsion, while tegus' tails are more cylindrical or even broader than high. In addition, tegus can run on their hind legs like a Collared Lizard, while monitors are quadrupedal.[18]

Economic importance and environmental impact

Tegus are among the most commercially exploited reptiles in the world. Up to 1,000,000 are harvested annually in their native Argentina for their hide and meat,[19] and are particularly important as a source of income in rural or indigenous communities.[20] Tegus can be also be found in captivity, where they are bred for the pet trade. They are reported to be highly intelligent, becoming docile as they mature and in some cases even ignoring food in favor of social interaction.[21] However, tegus have demanding husbandry requirements due to their large size.[22]

Within their native range, tegus are often thought of as pests, sometimes raiding chicken coops to feed on the eggs or fowl.[23] They are noted predators of ground nesting bird[24] and crocodilian eggs, and in some areas 80% of Spectacled Caiman nests are destroyed by tegus.[25] In South Florida, they have become an invasive species, and prey on the eggs of American Alligators instead. Predation by feral tegus may pose a threat to Florida's endangered wildlife, such as the Key Largo woodrat and the American Crocodile.[26] On account of their fruit eating habits, tegus may serve an important ecological function by dispersing seeds through their droppings.[27]

Taxonomy and evolution

Species listed alphabetically by specific name.[28]

The prominent pair of loreal scales on this black and white tegu identify it as belonging to the genus Salvator.

Mitochondrial DNA analysis indicates a deep divergence between a northern clade (containing T. teguixin, T. longilineus, T. palustris and T. quadrilineatus) and a southern clade (containing T. merianae, T. rufescens and T. duseni.).[30] The northern and southern clades are morphologically distinct, with the northern clade possessing a single pair of loreal scales between the eye and the nostril and a smooth texture to the scales on the body, and the southern clade possessing two pairs of loreal scales and a bumpy texture to the scales on the body.[31] At least one recent review of the morphology of the Teiidae family has placed the tegus of the southern clade in the genus Salvator.[32] Subsequent studies support the paraphyletic status of Tupinambis, though further research will be necessary to determine if the split will gain wider acceptance among the herpetological community.[33] Comparative analysis of hemipenial anatomy also provides support for the split between Tupinambis and Salvator.[34] Tegus probably originated sometime during the Cenozoic era. Tupinambis fossils from Argentina date back to the late-Miocene period.[35] Fossils of the extinct tegu Paradracaena can be found in earlier Miocene deposits.[36]

Name

As with many other animals from tropical South America (e.g. the Cariamae), the Tupinambis owes its scientific name to the pioneering accounts given by Piso & Marcgrave in their Historia Naturalis Brasiliae (1648). There was, however, a misinterpretation (by Linnaeus) of the Latin text, which reads "TEIVGVACV [...] Tupinambis", 'to the Tupinambá [Indians] TEIVGVACU'. Tupinambis was merely a metalinguistic term meaning 'to/for the Tupinambá,' whereas the intended, indigenous name for the animal was teiú-guaçú [lizard-big], lit. 'big lizard'.[37]

References

  1. ↑ Harvey, MB; Ugueto, GN; Gutberlet, RL (2012). "Review of Teiid Morphology with a Revised Taxonomy and Phylogeny of the Teiidae (Lepidosauria: Squamata)". Zootaxa. 3459: 1–156.
  2. ↑ http://www.zoologico.sp.gov.br/repteis/teiu.htm
  3. ↑ http://www.floridainvasives.org/Heartland/links/TeguBioprofileSep2006.pdf
  4. 1 2 3 4 5 6 7 8 9 Dessem, D (1985). "Ontogenetic Changes in the Dentition and Diet of Tupinambis (Lacertilia: Teiidae)". Copeia. 1985: 245–247. doi:10.2307/1444823.
  5. ↑ Sheffield, K. Megan; Butcher, Michael T.; Shugart, S. Katharine; Gander, Jennifer C.; Blob, Richard W. (2011). "Locomotor loading mechanics in the hindlimbs of tegu lizards (Tupinambis merianae): Comparative and evolutionary implications". The Journal of Experimental Biology. 214: 2616–2630. PMID 21753056. doi:10.1242/jeb.048801.
  6. ↑ http://www.myfwc.com/media/2380549/Tegu-brochure.pdf
  7. ↑ http://www.lvzoo.org/animal-profiles/tegu.cfm
  8. ↑ Rieppel, O. 1980. The Trigeminal Jaw Adductor Musculature of Tupinambis, with Comments on the Phylogenetic Relationships of the Teiidae (Reptilia, Lacertilia). Zool. J. Linne. Soc. Vol. 69(1):1–29.
  9. ↑ McBrayer, L.D., White, T.D. 2002. Bite Force, Behavior, and Electromyography in the Teiid Lizard, Tupinambis teguixin. Copeia. No. 1:111–119.
  10. 1 2 Pianka, E.R., Vitt, L.J. 2003. Lizards: Windows to the Evolution of Diversity. California. U. Cal. Press.
  11. ↑ Naretto, S.; Cardozo, G.; Blengini, C.S.; Chiaraviglio, M. (2014). "Sexual Selection and Dynamics of Jaw Muscle in Tupinambis Lizards". Evol. Biol. 41 (2): 192–200. doi:10.1007/s11692-013-9257-0.
  12. ↑ http://advances.sciencemag.org/content/2/1/e1500951 "Seasonal reproductive endothermy in tegu lizards"
  13. ↑ http://www.herpetologynotes.seh-herpetology.org/Volume6_PDFs/Sazima_HerpetologyNotes_volume6_pages427-430.pdf
  14. ↑ Kiefer, Mara Cíntia; Ivan Sazima (2002). "Diet of juvnile tegu lizard Tupinambis merianae (Teiidae) in southeastern Brazil" (PDF). Amphibia-Reptilia. 23: 105–108. Retrieved 24 October 2011.
  15. ↑ Hermano Del Duque. "Abundance of tegu lizards (Tupinambis merianae) in a remnant of the Brazilian Atlantic forest". Retrieved 27 March 2016.
  16. ↑ Aggressive Behavior and Performance in the Tegu Lizard Tupinambis merianae: Anthony Herrel, Denis V. Andrade, José Eduardo de Carvalho, Ananda Brito, Augusto Abe, and Carlos Navas - Physiological and Biochemical Zoology Vol. 82, No. 6 (November/December 2009), pp. 680-685
  17. ↑ Haddad, V; Duarte, MR; Neto, DG (2008). "Tegu (teiu) bite: report of human injury caused by a Teiidae lizard". Wilderness Environ Med. 19: 111–3. PMID 18513108. doi:10.1580/07-WEME-CR-1172.1.
  18. ↑ Monitors and Tegus, R. D. Bartlett and Patricia P. Bartlett
  19. ↑ http://agrilifecdn.tamu.edu/fitzgerald/files/2012/07/Mieres_Fitzgerald_2006_tegu-monitoring_7.pdf
  20. ↑ 1.Activity and Ranging Behavior of the Red Tegu Lizard Tupinambis rufescens in the Bolivian Chaco - Rossy R. Montaño, Rosa Leny Cuéllar, Lee A. Fitzgerald, Florencio Mendoza, Filemón Soria, Christine V. Fiorello, Sharon L. Deem, and Andrew J. Noss - South American Journal of Herpetology 2013 8 (2), 81-88 http://www.bioone.org/doi/abs/10.2994/SAJH-D-13-00016.1
  21. ↑ "Sisco Reptiles - Do tegu lizards seek human attention? Tupinambis Merianae Argentine Black & White Tegu, SiscoReptiles.com - The Original Tegu Attention Youtube Video". Archived from the original on 25 July 2014. Retrieved 27 March 2016.
  22. ↑ "Reptilecare.com - Argentine Black & White Tegus". Retrieved 27 March 2016.
  23. ↑ Guide to Lizards, Robert G. Sprackland, Ph.D.
  24. ↑ Mauro Galetti. "Density of the tegu lizard (Tupinambis merianae) and its role as nest predator at Anchieta island, Brazil". Retrieved 27 March 2016.
  25. ↑ "ADW: Caiman crocodilus: INFORMATION". Animal Diversity Web. Retrieved 27 March 2016.
  26. ↑ http://www.miamiherald.com/2013/09/30/3660515/hunt-is-on-for-tegu-lizards-in.html
  27. ↑ "Frugivory and seed dispersal by the tegu lizard Tupinambis merianae Reptilia: Teiidae - Castro - Papéis Avulsos de Zoologia (São Paulo)". Retrieved 27 March 2016.
  28. ↑ Tupinambis, The Reptile Database
  29. ↑ "Tegu - Tupinambis - Overview - Encyclopedia of Life". Encyclopedia of Life. Retrieved 27 March 2016.
  30. ↑ Fitzgerald et all. 1999
  31. ↑ "Salvator merianae". The Reptile Database. Retrieved 27 March 2016.
  32. ↑ Harvey, MB; Ugueto, GN; Gutberlet, RL (2012). "Review of teiid morphology with a revised taxonomy and phylogeny of the Teiidae (Lepidosauria: Squamata)" (PDF). Zootaxa. 3459: 1–156.
  33. ↑ Pyron, R. A.; Burbrink, F. T.; Wiens, J. J. (2013). "A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes". BMC Evolutionary Biology. 13 (1): 93. PMC 3682911 Freely accessible. PMID 23627680. doi:10.1186/1471-2148-13-93.
  34. ↑ Da Silva, M; Filho, G; Cronemberger, Á; Carvalho, L; Manzani, P; Vieira, J (2013). "Description of the hemipenial morphology of Tupinambis quadrilineatus Manzani and Abe, 1997 (Squamata, Teiidae) and new records from Piauí, Brazil". ZooKeys. 361: 61–72. doi:10.3897/zookeys.361.5738.
  35. ↑ Santiago Brizuela. "New Tupinambis remains from the late Miocene of Argentina and a review of the South American teiids". Retrieved 27 March 2016.
  36. ↑ Pujos, F.; Albino, A.M.; Baby, P.; Guyot, J.L. (2009). "Presence of the extinct lizard Paradracaena (Teiidae) in the Middle Miocene of the Peruvian Amazon". Journal of Vertebrate Paleontology. 29 (2): 594–598. doi:10.1671/039.029.0227.
  37. ↑ Cf. 'Etnolingüística' discussion list; 2/22/2012; http://lista.etnolinguistica.org/3167

Further reading

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