View source for Gopherus

{{Short description|Genus of tortoises}}
{{Automatic taxobox
| fossil_range = {{Fossil range|Priabonian|Recent}}
| image = Gopherus_polyphemus_Tomfriedel.jpg
| image_caption = Gopher tortoise, ''[[Gopherus polyphemus]]''
| taxon = Gopherus
| authority = [[Constantine Samuel Rafinesque-Schmaltz|Rafinesque]], 1832
| subdivision_ranks = Species
| subdivision =
* ''G. agassizii'' <small>(Cooper, 1863)</small>
* ''G. berlandieri'' <small>(Agassiz, 1857)</small>
* ''G. evgoodei'' <small>Edwards et al., 2016</small>
* ''G. flavomarginatus'' <small>Legler, 1959</small>
* ''G. morafkai'' <small>Murphy et al., 2011</small>
* ''G. polyphemus'' <small>(Daudin, 1802)</small>
}}
'''''Gopherus''''' is a [[genus]] of [[fossorial]] [[tortoise]]s commonly referred to as '''gopher tortoises'''. The gopher tortoise is grouped with land tortoises that originated 60 million years ago, in North America. A genetic study has shown that their closest relatives are in the [[Asia]]n genus ''[[Manouria]]''.<ref name = "Le et al.">{{cite journal  | last = Le  | first = M.  |author2=Raxworthy, C. J. |author3=McCord, W. P. |author4=Mertz, L.   | title = A molecular phylogeny of tortoises (Testudines: Testudinidae) based on mitochondrial and nuclear genes  | journal = Molecular Phylogenetics and Evolution  | volume = 40  | issue = 2  | pages = 517–531  | date = 2006-05-05  | doi = 10.1016/j.ympev.2006.03.003  | pmid = 16678445 }}</ref>
The gopher tortoises live in the [[southern United States]] from [[California]]'s [[Mojave Desert]] across to [[Florida]], and in parts of northern [[Mexico]]. Gopher tortoises are so named because of some species' habit of digging large, deep burrows ([[gopher]]s are small terrestrial burrowing rodents). Most notably, ''[[Gopherus polyphemus]]'' digs burrows which can be up to {{convert|40|ft|m}} in length and {{convert|10|ft|m}} in depth.<ref>{{cite web|url=http://www.gophertortoisecouncil.org/about.php |title=About The Gopher tortoise |publisher=Gopher Tortoise Council |access-date=2009-07-03 |archive-url=https://web.archive.org/web/20090901200608/http://www.gophertortoisecouncil.org/about.php |archive-date=2009-09-01 }}</ref> These burrows are used by a variety of other species, including mammals, other reptiles, amphibians, and birds.<ref name="Turtles">{{cite book |url=https://books.google.com/books?id=CQ4Trk0_K5wC |title=Turtles |first=Charles |last=Fergus |publisher=Stackpole Books |year=2007 |isbn=978-0-8117-3420-2 |pages=22–24}}</ref>  Gopher tortoises are {{convert|20|-|50|cm|in|abbr=on}} in length, depending on the species.<ref name="Franklin">{{cite book |url=https://books.google.com/books?id=rBbuYFjiTNMC |title=Turtles: An Extraordinary Natural History 245 Million Years in the Making |first=Carl J. |last=Franklin |publisher=MBI Publishing Company |year=2007 |isbn=978-0-7603-2981-8 |pages=129–131}}</ref>  All six species are found in [[xeric]] habitats. Numerous extinct species are known, the oldest dating to the [[Priabonian]] stage of the Late Eocene of the United States.<ref>{{Cite web|url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=37741|title=Fossilworks: ''Gopherus''|website=fossilworks.org|access-date=2020-02-29}}</ref>
==Species==
[[File:Geographic distribution of the North American tortoises of the genus Gopherus.jpg|thumb|400 px|Geographic distribution of species within the genus ''Gopherus''|center]]In July 2011, researchers decided on the basis of DNA, and morphological and behavioral data that the Sonoran and Mojave populations of the desert tortoise, ''G. agassizii'' were distinct species.<ref name="usgs.gov">[http://www.usgs.gov/newsroom/article.asp?ID=2842&from=rss_home US Geological Survey]</ref> This newly described species was named ''G. morafkai'', or the Morafka's desert tortoise. The acceptance of ''G. morafkai'' reduced the range of ''G. agassizii'' by about 70%<ref>{{Cite journal|last1=Murphy|first1=Robert|last2=Berry|first2=Kristin|last3=Edwards|first3=Taylor|last4=Leviton|first4=Alan|last5=Lathrop|first5=Amy|last6=Riedle|first6=J. Daren|date=2011-06-28|title=The dazed and confused identity of Agassiz's land tortoise, Gopherus agassizii (Testudines: Testudinidae) with the description of a new species and its consequences for conservation|url=https://zookeys.pensoft.net/article/2586/|journal=ZooKeys|language=en|issue=113|pages=39–71|doi=10.3897/zookeys.113.1353|issn=1313-2970|pmc=3187627|pmid=21976992|doi-access=free|bibcode=2011ZooK..113...39M }}</ref>  In 2016, based on a large-scale genetic analysis, ecological and morphological data, researchers proposed a split between the Sonoran and Sinaloan populations.<ref name=":1" /> This southernmost member of the ''Gopherus'' genus was named ''G. evgoodei''.<ref name=":1" /> As such, there are currently six recognized extant species in the genus ''Gopherus'':
=== Extant ===
Listed alphabetically by binomial name:<ref>[http://reptile-database.reptarium.cz/advanced_search?genus=Gopherus ''Gopherus''], The Reptile Database</ref>
{| class="wikitable"
|-
! Image !! Common Name !! Scientific name !! Distribution
|-
| [[File:Desert tortoise.jpg|150x150px]]||Mojave desert tortoise || ''[[Gopherus agassizii]]''
<small>([[James Graham Cooper|Cooper]], 1863)</small>
|Endemic to USA. [[Mojave Desert]] in southeastern California,
southern Nevada, and extreme southwestern Utah and
northwest Arizona, north and west of the Colorado River.<ref name="Bramble & Hutchison (2014)">Bramble, Dennis M. and J. Howard Hutchison. 2014. ''Morphology, Taxonomy, and Distribution of North American Tortoises: An Evolutionary Perspective.'' pages 1-12 '''In''', David C. Rostal, Earl D. McCoy, and Henry R. Mushinsky (editors). ''Biology and Conservation of North American Tortoises.'' Johns Hopkins University Press. Baltimore, Maryland. x, 190 pp.{{ISBN|1-4214-1377-9}}</ref>{{rp|3–4 p.}}
|-
| [[File:Texas Tortoise (Gopherus berlandieri) (57391706).jpg|150x150px]]||Texas tortoise || ''[[Gopherus berlandieri]]''
<small>([[Louis Agassiz|Agassiz]], 1857)</small>
|[[Tamaulipan mezquital]] ecoregion of south Texas,
USA, eastern Coahuila, northern Nuevo León, and Tamaulipas,
Mexico with a few old records (questionable) from southern San Luis Potosí.<ref name="Rose and Judd (2014)">Rose, Francis L and Frank W. Judd. 2014. ''The Texas Tortoise: A Natural History.'' University of Oklahoma Press, Norman, Oklahoma. xviii, 188 pp.{{ISBN|978-0-8061-4451-1}}</ref>{{rp| 25 & 27p.}}
|-
| [[File:Gopherus evgoodei, Sonora, Mexico.jpg|150x150px]]|| Goode's thornscrub tortoise || ''[[Gopherus evgoodei]]'' <small>Edwards et al., 2016</small><ref name=":1">{{cite journal|vauthors=Edwards T, Karl AE, Vaughn M, Rosen PC, Torres CM, Murphy RW|title=The desert tortoise trichotomy: Mexico hosts a third, new sister-species of tortoise in the ''Gopherus morafkai''–''G. agassizii'' group|journal=ZooKeys|date=2016|issue=562|pages=131–158|doi=10.3897/zookeys.562.6124|url=https://zookeys.pensoft.net/articles.php?id=6124&display_type=list&element_type=9|pmid=27006625|pmc=4768471|doi-access=free|bibcode=2016ZooK..562..131E }}</ref>||Endemic to Mexico. Tropical deciduous forest and thornscrub in southern Sonora and adjacent areas of extreme northern Sinaloa
and extreme southwestern Chihuahua.
|-
| [[File:Itzelbolson1small.jpg|150x150px]]||Bolson tortoise || ''[[Gopherus flavomarginatus]]''
<small>[[John M. Legler|Legler]], 1959</small>
|Endemic to Mexico. [[Chihuahuan Desert|Chihuahua Desert]] of North-central Mexico,
in the vicinity of the conjunction of the Chihuahua, Coahuila, and Durango state borders.
|-
| [[File:Sonoran Desert Tortoise FWS 11909.jpg|150x150px]] || Sonoran desert tortoise || ''[[Gopherus morafkai]]''
<small>Murphy et al., 2011</small><ref name="usgs.gov" />
| [[Sonoran Desert|Sonora Desert]] in northern Sonora, including Tiburon Island,
Mexico and southwest Arizona, south and east of the Colorado River.<ref name="Bramble & Hutchison (2014)" />{{rp|4 p.}}
|-
| [[File:Gopher Tortoise at Smyrna Dunes Park - Flickr - Andrea Westmoreland (1).jpg|150x150px]] || Gopher tortoise || ''[[Gopherus polyphemus]]''
<small>([[François Marie Daudin|Daudin]], 1802)</small>
|Endemic to USA. Extreme eastern Louisiana,
southern Mississippi, southern Alabama, southern Georgia,
Florida, and extreme southeastern South Carolina.
|-
|}
=== Fossil ===
Members of ''Gopherus'' that became extinct in the late [[Pleistocene]]:
* {{extinct}}''Gopherus donlaloi'' <small>(Reynoso and Montellano-Ballesteros, 2004)</small><ref>{{Cite journal|title = A New Giant Turtle of the Genus ''Gopherus'' (Chelonia: Testudinidae) from the Pleistocene of Tamaulipas, México, and a Review of the Phylogeny and Biogeography of Gopher Tortoises|jstor = 4524778|journal = Journal of Vertebrate Paleontology|date = 2004-01-01|pages = 822–837|volume = 24|issue = 4|first1 = Víctor-Hugo|last1 = Reynoso|first2 = Marisol|last2 = Montellano-Ballesteros|doi=10.1671/0272-4634(2004)024[0822:angtot]2.0.co;2| s2cid=84397134 | url=http://doc.rero.ch/record/15347/files/PAL_E2686.pdf }}</ref><ref>{{Cite book|title = Biology and Conservation of North American Tortoises|url = https://books.google.com/books?id=CKr6AwAAQBAJ|publisher = JHU Press|date = 2014-06-13|isbn = 9781421413778|language = en|first1 = David C.|last1 = Rostal|first2 = Earl D.|last2 = McCoy|first3 = Henry R.|last3 = Mushinsky}}</ref><ref>{{Cite web|url = http://www.torreyaguardians.org/hansen-2010.pdf|title = Ecological history and latent conservation potential: large and giant tortoises as a model for taxon substitutions}}</ref>
* {{extinct}}''Gopherus depressus''<ref name=Woodburne1972>Woodburne & Golz, 1972</ref>
* {{extinct}}''Gopherus hexagonatus'' (<small>Cope, 1893</small>) Late Pliocene-Late Pleistocene, large sized species, with carapaces over a metre (3.3&nbsp;ft) in length.<ref>{{Cite journal|last1=Esker|first1=Donald A.|last2=Forman|first2=Steve L.|last3=Butler|first3=Dava K.|date=May 2019|title=Reconstructing the mass and thermal ecology of North American Pleistocene tortoises|url=https://www.cambridge.org/core/product/identifier/S009483731900006X/type/journal_article|journal=Paleobiology|language=en|volume=45|issue=2|pages=363–377|doi=10.1017/pab.2019.6|bibcode=2019Pbio...45..363E |s2cid=155216574 |issn=0094-8373}}</ref>
==Breeding==
Gopher tortoises usually mate during April and May. The female will then choose either a sunny spot nearby or a sandy mound in front of her burrow to lay between 3 and 15 eggs. The eggs then hatch from 70 to 100 days later. Once hatched, the baby tortoises spend most of their time in their mother's burrow until they learn to dig their own burrow. They do not reach maturity until they are around 10 to 15 years old. Gopher tortoises have an abbreviated mating season in early spring, when male tortoises visit the female tortoise' burrows and mate with them.<ref>Landers, L. J.; Garner, J. A. & McRae, W. A. (1980). "Reproduction on Gopher Tortoises in Southwestern Georgia". ''Herpetological''. '''36''': 353–361.</ref>
==Diet==
Gopher tortoises are mainly [[herbivore]]s that feed on low growing plant life. Their diet consists mostly of [[grass]]es and [[legume]]s but they will also feed on small [[Berry|berries]] and [[fruit]]s.
[[File:Desert tortoise (Gopherus agassizii) drinking from roadway.jpg|thumb|250 px|Desert tortoise (''Gopherus agassizii'') drinking from roadway in [[Joshua Tree National Park]]]]
The diet of tortoises contain excess salt, sodium, chloride, and potassium that must be purged from the body, and drinking free standing water, even if only once or a few times each year, is essential for this function and for tortoise survival. Opportunities for gopher tortoises to drink water vary greatly between the species in the genus. ''Gopherus agassizii'' live in extremely arid areas that can receive as little as 10–20&nbsp;cm. per year. ''Gopherus polyphemus'' live in [[mesic habitat]], where water is availability and evaporative loss is less problematic. Other species live in intermediate environments where a few weeks of rain typically occur twice per year, or where relatively consistent summer rains occur. When pools of rain water or saturated soils are available tortoises will direct their heads face down and submerge their face to a level just below the eyes and drink copiously. Reptiles modify urine and plasma concentrations in their bodies with their bladder, cloaca, and colon, rather than the kidneys. The bladder plays a major function in regulating blood osmolality: permeable to ammonia, urea, water, and small ions, but not uric acid. This permeability allows tortoises that are hibernating, or living in arid environments without drinking water for months at a time to store uric acid, but resorb water from the bladder. Tortoises react to dry periods by retreating to shelters (burrows, caves etc.) with more humid [[microhabitats]], and remaining inactive. Gopher tortoises can survive a year of drought through both behavior and physiological adaptations, two years of drought can result in deteriorating body conditions, and extended years of drought will produce high mortalities of gopher tortoises.<ref name="Esque (2014)">Esque, Todd, Kristina Drake, and Kenneth Nussear. 2014. ''Water and Food Acquisition and Their Consequences for Life History and Metabolism of North American Tortoises.'' pages 85-95. In, David C. Rostal, Earl D. McCoy, and Henry R. Mushinsky (editors). ''Biology and Conservation of North American Tortoises.'' Johns Hopkins University Press. Baltimore, Maryland. x, 190 pp.{{ISBN|1-4214-1377-9}}</ref>{{rp|85–87, 94 pp.}}
There are many observations of ''Gopherus'' eating non-vegetation food items. Documented examples include a variety of bones, snail shells, soil at mineral licks, charcoal, sand, stones, human trash, carrion, raptor pellets and various animals feces. ''Gopherus polyphemus'' studies observed specimens moving bones into their burrows and found the fourth most common matter in their scats was insect material. The exact reasons are not entirely understood, some cases might simply be due to accidentally ingesting materials near food items or sampling potential foods. Hypotheses concerning the consumption of animal matter by ''Gopherus'' include supplements for the low levels of calcium, phosphorus, or protein in their diet, and a need for [[calcium carbonate]] in bones, the production of eggshells in females, and growing young. Hypotheses on the consumption of nonfood items such as soil or rocks include [[mastication]] or [[vermifuge]] for the removal of parasites.<ref name="Esque (2014)" />{{rp|93 p.}}
==Conservation==
[[File:Gopherus berlandieri, Texas Tortoise, Tamaulipas.jpg|thumb|250 px|left|[[Texas tortoise]] (''Gopherus berlandieri''), northern [[Tamaulipas]], Mexico]]
Populations of all six species of ''Gopherus'' have declined dramatically. In the past, gopher tortoises were hunted for their meat, which was used in [[Turtle soup|stews]].<ref>"Gopher Tortoise Stew", in: [http://www.smithsonianmag.com/people-places/journeys_recipe.html?c=y&page=2 Recipes from Another Time: Savor the flavor of old St. Augustine and try a couple of these original recipes].  Smithsonian magazine, October 2001</ref> Currently the most significant threat to their survival is [[habitat destruction]], but the pet trade and collisions with vehicles have also taken their toll.<ref name="Franklin"/> To help decrease gopher tortoise death due to collisions with vehicles, the US Department of Transportation in Mississippi has recently placed angled fences along the road side to keep tortoise from wandering onto the highways near their habitats.<ref>{{cite web |title= Fence with an "angle" protects gopher tortoises |url=http://www.fhwa.dot.gov/environment/wildlifeprotection/index.cfm?fuseaction=home.viewArticle&articleID=44|archive-url=https://web.archive.org/web/20030625154517/http://www.fhwa.dot.gov/environment/wildlifeprotection/index.cfm?fuseaction=home.viewArticle&articleID=44|url-status=dead|archive-date=June 25, 2003| publisher=US Department of Transportation | date = 2003-04-25 | access-date = 2010-04-18}}</ref> On November 9, 2009, the US Fish and Wildlife Service proposed rulemaking to include the eastern population of the gopher tortoise, ''Gopherus polyphemus'', in the List of Threatened Wildlife.<ref>{{cite periodical |url=http://www.fws.gov/policy/library/2009/E9-26841.pdf |periodical=Federal Register |title=Department of the Interior: Fish and Wildlife Service: 50 CFR Part 17 | volume = 74| issue = 215 | date = 2009-11-09 |access-date=2009-11-13}}</ref> In 2018, the IUCN Tortoise and Freshwater Turtle Specialist Group recommended a re-assessment and re-classification of all six ''Gopherus'' species<ref name=":0">{{Cite journal|last1=Rhodin|first1=Anders G. J.|last2=Stanford|first2=Craig B.|last3=Dijk|first3=Peter Paul Van|last4=Eisemberg|first4=Carla|last5=Luiselli|first5=Luca|last6=Mittermeier|first6=Russell A.|last7=Hudson|first7=Rick|last8=Horne|first8=Brian D.|last9=Goode|first9=Eric V.|last10=Kuchling|first10=Gerald|last11=Walde|first11=Andrew |date= December 2018 |title=Global Conservation Status of Turtles and Tortoises (Order Testudines)|url=https://bioone.org/journals/chelonian-conservation-and-biology/volume-17/issue-2/CCB-1348.1/Global-Conservation-Status-of-Turtles-and-Tortoises-Order-Testudines/10.2744/CCB-1348.1.full|journal=Chelonian Conservation and Biology|volume=17|issue=2|pages=135–161|doi=10.2744/CCB-1348.1|s2cid=91937716|issn=1071-8443}}</ref> This reclassification would move ''G. agassizii'' from Vulnerable (VU) to Critically Endangered (CR), ''G. berlandieri'' from Near Endangered (NE) to Near Threatened (NT), ''G. evgoodei'' from Near Endangered (NE) to Vulnerable (VU), ''G. flavomarginatus'' from Vulnerable (VU) to Critically Endangered (CR), ''G. morafkai'' from Near Endangered (NE) to Vulnerable (VU) and ''G. polyphemus'' from Vulnerable (VU) to Endangered (EN).<ref name=":0" />
==Gallery==
<gallery mode="packed" heights="150px">
File:Gopherus flavomarginatus 403375 (cropped).jpg|A juvenile bolson tortoise (''Gopherus flavomarginatus''), from Durango, Mexico (7 July 2006)
File:DesertTortoise.JPG|Desert tortoise (''[[Gopherus agassizii]]'') [[San Bernardino County, California|San Bernardino County]], California, US.
File:Goph, Gopherus polyphemus, in Sanibel Island, Florida 01 (cropped).jpg|Gopher tortoise (''Gopherus polyphemus'') from Lee County, Florida (18 June 2018)
File:Big Boy ( Gopherus Polyphemus from New Smyrna Beach, Florida ) - Flickr - Andrea Westmoreland.jpg|Gopher tortoise (''Gopherus polyphemus'') from Volusia County, Florida (14 September 2008)
</gallery>
{{clear left}}
== References ==
{{Reflist}}
=== Bibliography ===
* {{cite book |last1=Woodburne |first1=M. O. |first2=D. J. |last2=Golz |year=1972 |title=Stratigraphy of the Punchbowl Formation, Cajon Valley, southern California |series=[[University of California Publications in Geological Sciences]] |volume=92}}
==External links==
{{Commons category}}
{{Wikispecies}}
* [http://www.gophertortoisecouncil.org Gopher Tortoise Council]
* [http://www.gophertortoise.org The Gopher Tortoise Organization]
{{Testudinidae}}
{{Taxonbar|from=Q1012045}}
{{Authority control}}
[[Category:Gopherus| ]]
[[Category:Turtle genera]]
[[Category:Taxa named by Constantine Samuel Rafinesque]]