Heloderma

The genus Heloderma contains the Gila monster (H. suspectum) and four species of beaded lizards. Their eyes are immobile and fixed in their heads.{{Cite book|url=https://books.google.com/books?id=ORptEAAAQBAJ&dq=Extraocular+muscles+Heloderma&pg=PA206|title=Wild and Exotic Animal Ophthalmology: Volume 1: Invertebrates, Fishes, Amphibians, Reptiles, and Birds|first1=Fabiano|last1=Montiani-Ferreira|first2=Bret A.|last2=Moore|first3=Gil|last3=Ben-Shlomo|date=April 27, 2022|publisher=Springer Nature|isbn=978-3-030-71302-7 |via=Google Books}}{{Cite book|url=https://books.google.com/books?id=SuKRyC1Q8W4C&dq=Gila+monster+eyes+fixed+in+their+head&pg=PA8|title=The Biology of the Eye|first=Jorge|last=Fischbarg|date=December 9, 2005|publisher=Elsevier|isbn=978-0-08-047609-4 |via=Google Books}} The Gila monster is a large, stocky, mostly slow-moving reptile that prefers arid deserts. Beaded lizards are seen to be more agile and seem to prefer more humid surroundings.{{Cite journal|last=C. M. Bogert, R. M. Del Campo|date=1956|title=The Gila Monster and its Allies. The relationships, habits, and behavior of the lizards of the family Helodermatidae|url=|journal=Bulletin of the American Museum of Natural History|volume=109|pages=1–238|via=}}{{Cite book|last=Beck|first=D. D.|title=Biology of Gila Monsters and Beaded Lizards|publisher=University Press of California|year=2005|isbn=|location=|pages=}} The tails of all species of Heloderma are used as fat-storage organs. The scales of the head, back, and tail are bead-like, containing osteoderms for better protection. The scales of the belly are free from osteoderms. Most species are dark in color, with yellowish or pinkish markings.{{Cite book|last=Schwandt|first=Hans-Joachim|title=The Gila Monster Heloderma suspectum|publisher=Edition Chimaira|year=2019|isbn=978-3-89973-441-6|location=Frankfurt/Main|pages=}}{{cite book |editor1=Cogger, H.G. |editor2=Zweifel, R.G. |author = Bauer, Aaron M.|year=1998|title=Encyclopedia of Reptiles and Amphibians|publisher= Academic Press|location=San Diego|pages= 156|isbn= 0-12-178560-2}}

The venom glands of Heloderma are located at the end of the lower jaws, unlike snakes' venom glands, which are located behind the eyes. Also, unlike snakes, the Gila monster and beaded lizards lack the musculature to inject venom immediately. They have to chew the venom into the flesh of a victim. Heloderma venom is used only in defense. Venom glands are believed to have evolved early in the lineage leading to the modern helodermatids, as their presence is indicated even in the 65-million-year-old fossil genus Paraderma.Richard L. Cifelli, Randall L. Nydam. 1995. Primitive, helodermatid-like platynotans from the Early cretaceous of Utah. Herpetologica. 51(3):286-291. In general, one adult helodermatid has around 15 to 20 mg of venom, while the estimated lethal dose for humans is 5 to 8 mg.{{Cite book|last=Dart|first=Richard C.|url=https://books.google.com/books?id=BfdighlyGiwC&dq=gila+monster+estimated+lethal+dose+for+human&pg=PA1539|title=Medical Toxicology|date=2004|publisher=Lippincott Williams & Wilkins|isbn=978-0-7817-2845-4|language=en}}

Venom production among lizards was long thought to be unique to this genus, but researchers studying venom production have proposed many others also produce some venom, all placed in the clade Toxicofera, which includes all snakes and 13 other families of lizards..

{{cite journal |author=Fry, B. |display-authors=etal |date=February 2006 |title=Early evolution of the venom system in lizards and snakes |journal=Nature |volume=439 |issue=7076 |pages=584–588 |bibcode=2006Natur.439..584F |doi=10.1038/nature04328 |pmid=16292255 |s2cid=4386245}} However, except for snakes, helodermatids, and possibly varanids, envenomation is not considered medically significant for humans.

Helodermatids are carnivorous, preying on rodents and other small mammals, and eating the eggs of birds and reptiles.

All species of Heloderma are oviparous. The Gila monster typically lays six eggs, and the beaded lizards up to about 18 eggs . Comparing the different species, all eggs have a similar size, as are their hatchlings.

[[File:Kladoramm heloderma.jpg|alt=Cladogram of the heloderma species|thumb|Cladogram of the Heloderma species

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Family Helodermatidae

The four subspecies of beaded lizards were elevated to full species in 2013.http://www.redlist-arc.org/Article-PDFs/Special%20Mexico%20Issue_ARC_7(1)_74-96_low_res.pdf {{Webarchive|url=https://web.archive.org/web/20131023060725/http://www.redlist-arc.org/Article-PDFs/Special%20Mexico%20Issue_ARC_7%281%29_74-96_low_res.pdf |date=2013-10-23 }} Reiserer & al., 2013, Taxonomic reassessment and conservation status of the beaded lizard, Heloderma horridum (Squamata: Helodermatidae)

File:Fossil Florida Heloderma FLMNH.jpg of Florida (FLMNH). These lizards are no longer found in the region.]]

Members of the genus Heloderma have many extinct relatives in the Helodermatidae, whose evolutionary history may be traced back to the Cretaceous period, such as Estesia. The genus Heloderma has existed since the Miocene, when H. texana lived, and fragments of osteoderms from the Gila monster have been found in late Pleistocene (8,000-10,000 years ago) deposits near Las Vegas, Nevada. Because the helodermatids have remained relatively unchanged morphologically, they are occasionally regarded as living fossils.{{cite book |author1=King, Ruth Allen |author2=Pianka, Eric R. |author3=King, Dennis |title=Varanoid Lizards of the World |publisher=Indiana University Press |location=Bloomington |year=2004 |isbn=0-253-34366-6 }} Although the beaded lizards and the Gila monster appear closely related to the monitor lizards (varanids) of Africa, Asia, and Australia, the wide geographical separation and unique features not found in the varanids indicate they are better placed in a separate family.{{cite book |author=Mattison, Chris |title=Lizards of the World |publisher=Blandford |location=London |year=1998 |isbn=0-7137-2357-2 |url-access=registration |url=https://archive.org/details/lizardsofworld00matt }}

File:Beaded Lizard Skeleton.jpg)|left]]

The type species is Heloderma horridum, which was first described in 1829 by Arend Wiegmann. Although he originally assigned it the generic name Trachyderma, he changed it to Heloderma six months later, which means "studded skin", from the Ancient Greek words hêlos (ηλος)—the head of a nail or stud—and derma (δερμα), meaning skin.{{cite journal | last = Wiegmann| first = A.F.A.| author-link = Arend Friedrich August Wiegmann| title = Über die Gesetzlichkeit in der geographischen Verbreitung der Saurier| journal = Isis| volume =22 | issue =3–4 | pages = 418–428| publisher =Oken | year = 1829 }}

Conrad, 2008 and Estes et al., 1988 (using morphological data) place the Helodermatidae within Varanoidea along with Lanthanotus borneensis and Varanus.{{Cite journal|last1=Conrad|first1=Jack L.|last2=Ast|first2=Jennifer C.|last3=Montanari|first3=Shaena|last4=Norell|first4=Mark A.|date=2011|title=A combined evidence phylogenetic analysis of Anguimorpha (Reptilia: Squamata)|journal=Cladistics|language=en|volume=27|issue=3|pages=230–277|doi=10.1111/j.1096-0031.2010.00330.x|pmid=34875778|s2cid=84301257|issn=1096-0031|doi-access=free}}{{Cite journal|last=Estes|first=Richard|date=1988|title=Phylogenetic relationships within squamata|url=https://repository.si.edu/handle/10088/6457|journal=Phylogenetic Relationships of the Lizard Families: Essays Commemorating Charles L. Camp|pages=119–281|hdl=10088/6457|isbn=9780804714358}} However, Estes et al., 1988 understood the Helodermatidae as having split earlier from Lanthanotus and Varanus, whereas Conrad, 2008 groups them at the same branch point.

In contrast, molecular studies have identified Heloderma as being within the Anguioidea along with Anguidae and Xenosauridae, but specifically sister to Anguidae.{{Cite journal|last1=Vidal|first1=Nicolas|last2=Marin|first2=Julie|last3=Sassi|first3=Julia|last4=Battistuzzi|first4=Fabia U.|last5=Donnellan|first5=Steve|last6=Fitch|first6=Alison J.|last7=Fry|first7=Bryan G.|last8=Vonk|first8=Freek J.|last9=Rodriguez de la Vega|first9=Ricardo C.|last10=Couloux|first10=Arnaud|last11=Hedges|first11=S. Blair|date=2012-10-23|title=Molecular evidence for an Asian origin of monitor lizards followed by Tertiary dispersals to Africa and Australasia|url= |journal=Biology Letters|volume=8|issue=5|pages=853–855|doi=10.1098/rsbl.2012.0460|pmc=3441001|pmid=22809723}}{{Cite journal|last1=Townsend|first1=Ted M.|last2=Larson|first2=Allan|last3=Louis|first3=Edward|last4=Macey|first4=J. Robert|date=2004-10-01|title=Molecular Phylogenetics of Squamata: The Position of Snakes, Amphisbaenians, and Dibamids, and the Root of the Squamate Tree|journal=Systematic Biology|language=en|volume=53|issue=5|pages=735–757|doi=10.1080/10635150490522340|pmid=15545252|issn=1063-5157|doi-access=free}}File:Heloderma suspectum skull with dentition.jpg

File:Heloderma hatchling sizes.jpg

H. horridum, H. exasperatum, and H. suspectum are frequently found in captivity and are well represented in zoos throughout much of the world. The other two species of Heloderma, H. alvarezi and H. charlesbogerti, are extremely rare, and only a few captive specimens are known.

File:Heloderma Eiablage.jpg|alt=Helderema suspectum with 4 eggs|Heloderma suspectum with four eggs

File:Gila-monster-6-eggs.jpg|alt=Helderema suspectum with 6 eggs|Heloderma suspectum with six eggs

File:Schluepfendes-jungtier-OS6.jpg|alt=Gila monster hatching|Gila monster hatching

File:Group of young gila monster.jpg|alt=Group of young Gila monsters|Group of young Gila monsters

File:Heloderma Skin.jpg

File:Heloderma suspectum captured.jpg

File:H-horridum.jpg

{{Reflist}}Image:Heloderma_alvarezi2.jpg

• {{cite journal|last1=Ariano-Sánchez|first1=Daniel|title=Envenomation by a wild Guatemalan beaded lizard Heloderma horridum charlesbogerti|url=|journal=Clinical Toxicology|date=2008|volume=46|issue=9|pages=897–899|doi=10.1080/15563650701733031|pmid=18608297|s2cid=22173811|doi-access=free}}
• Ariano-Sánchez, D. & G. Salazar. 2007. Notes on the distribution of the endangered lizard, Heloderma horridum charlesbogerti, in the dry forests of eastern Guatemala: an application of multi-criteria evaluation to conservation. Iguana 14: 152–158. [https://www.scribd.com/doc/11116925/Ariano-D-y-G-Salazar-2007-Notes-on-the-Distribution-of-the-Endangered-Lizard-Heloderma-horridum-charlesbogerti-in-the-Dry-Forests-of-Eastern-Gu]
• Ariano-Sánchez, D. 2006. The Guatemalan beaded lizard: endangered inhabitant of a unique ecosystem. Iguana 13: 178–183. [https://www.scribd.com/doc/11117893/Ariano-D-2006-The-Guatemalan-beaded-lizard-Endangered-inhabitant-of-a-unique-ecosystem-Iguana-133-178183]
• [http://www.ssn.org/Meetings/cop/cop14/Other/CoP14_results.pdf CONVENTION ON INTERNATIONAL TRADE IN ENDANGERED SPECIES OF WILD FAUNA AND FLORA] {{Webarchive|url=https://web.archive.org/web/20180930041659/http://www.ssn.org/Meetings/cop/cop14/Other/CoP14_results.pdf |date=2018-09-30 }}. 2007. Resume of the 14th Convention of the Parts. The Hague. The Netherlands.

File:Heloderma exasperatum.jpg

{{Wikispecies|Helodermatidae}}

Schwandt, Hans- Joachim [https://www.heloderma.net/en/ www.heloderma.net] 2006 in 6 languages

{{Commons category|Heloderma}}

• {{Cite book|last=C. M. Bogert, R. M. Del Campo|title=The Gila Monster and its Allies. The relationships, habits, and behavior of the lizards of the family Helodermatidae|publisher=Bulletin of the American Museum of Natural History|year=1956|isbn=|volume=109|location=|pages=1–238}}
• {{Cite book|last=Beck|first=D. D.|title=Biology of Gila Monsters and Beaded Lizards|publisher=University Press of California|year=2005|isbn=|location=London|pages=}}
• {{Cite book |last=Schwandt |first=Hans-Joachim |title=The Gila Monster Heloderma suspectum - Natural History, Husbandry & Propagation |publisher=Edition Chimaira |year=2019 |isbn=978-3-89973-441-6 |location=Frankfurt/Main |pages=}}

• [http://www.helodermahorridum.com/beaded_lizard.php About Beaded Lizards] {{Webarchive|url=https://web.archive.org/web/20150215024644/http://www.helodermahorridum.com/beaded_lizard.php |date=2015-02-15 }}
• [http://herpetology.com/heloderma.html Heloderma information]
• [http://www.jcvi.org/reptiles/families/helodermatidae.php Family Helodermatidae (Gila Monsters)] {{Webarchive|url=https://web.archive.org/web/20100715164854/http://www.jcvi.org/reptiles/families/helodermatidae.php |date=2010-07-15 }}

{{Squamata families}}

{{Varanoidea}}

{{Taxonbar|from=Q729412}}

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Category:Extant Miocene first appearances

Category:Lizard genera

Category:Taxa named by Arend Friedrich August Wiegmann