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Gopherus

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Genus of tortoises

Gopherus Temporal range:Priabonian–Recent PreꞒ Ꞓ O S D C P T J K Pg N

Gopher tortoise,Gopherus polyphemus
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Order:	Testudines
Suborder:	Cryptodira
Family:	Testudinidae
Genus:	Gopherus Rafinesque, 1832
Species
Gopherus agassizii(Cooper, 1863) Gopherus berlandieri(Agassiz, 1857) Gopherus evgoodeiEdwards et al., 2016 Gopherus flavomarginatusLegler, 1959 Gopherus morafkaiMurphy et al., 2011 Gopherus polyphemus(Daudin, 1802)

Distribution of the genusGopherus

Gopherus is agenus offossorial tortoises commonly referred to asgopher 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 theAsian genusManouria.^[1] The gopher tortoises live in thesouthern United States fromCalifornia'sMojave Desert across toFlorida, and in parts of northernMexico. Gopher tortoises are so named because of some species' habit of digging large, deep burrows (gophers are small terrestrial burrowing rodents). Most notably,Gopherus polyphemus digs burrows which can be up to 40 feet (12 m) in length and 10 feet (3.0 m) in depth.^[2] These burrows are used by a variety of other species, including mammals, other reptiles, amphibians, and birds.^[3] Gopher tortoises are 20–50 cm (7.9–19.7 in) in length, depending on the species.^[4] All six species are found inxeric habitats. Numerous extinct species are known, the oldest dating to thePriabonian stage of the Late Eocene of the United States.^[5]

Species

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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.^[6] This newly described species was namedG. morafkai, or the Morafka's desert tortoise. The acceptance ofG. morafkai reduced the range ofG. agassizii by about 70%^[7] In 2016, based on a large-scale genetic analysis, ecological and morphological data, researchers proposed a split between the Sonoran and Sinaloan populations.^[8] This southernmost member of theGopherus genus was namedG. evgoodei.^[8] As such, there are currently six recognized extant species in the genusGopherus:

Extant

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Listed alphabetically by binomial name:^[9]

Common name	Scientific name	Distribution
Mojave desert tortoise	Gopherus agassizii (Cooper, 1863)	Endemic to USA.Mojave Desert in southeastern California, southern Nevada, and extreme southwestern Utah and northwest Arizona, north and west of the Colorado River.^[10]^{: 3–4 p.}
Texas tortoise	Gopherus berlandieri (Agassiz, 1857)	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.^[11]^{: 25 & 27p.}
Goode's thornscrub tortoise	Gopherus evgoodeiEdwards et al., 2016^[8]	Endemic to Mexico. Tropical deciduous forest and thornscrub in southern Sonora and adjacent areas of extreme northern Sinaloa and extreme southwestern Chihuahua.
Bolson tortoise	Gopherus flavomarginatus Legler, 1959	Endemic to Mexico.Chihuahua Desert of North-central Mexico, in the vicinity of the conjunction of the Chihuahua, Coahuila, and Durango state borders.
Sonoran desert tortoise	Gopherus morafkai Murphy et al., 2011^[6]	Sonora Desert in northern Sonora, including Tiburon Island, Mexico and southwest Arizona, south and east of the Colorado River.^[10]^{: 4 p.}
Gopher tortoise	Gopherus polyphemus (Daudin, 1802)	Endemic to USA. Extreme eastern Louisiana, southern Mississippi, southern Alabama, southern Georgia, Florida, and extreme southeastern South Carolina.

Fossil

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Members ofGopherus that became extinct in the latePleistocene:

†Gopherus donlaloi(Reynoso and Montellano-Ballesteros, 2004)[12]^[13]^[14]
†Gopherus depressus[15]
†Gopherus hexagonatus (Cope, 1893) Late Pliocene-Late Pleistocene, large sized species, with carapaces over a metre (3.3 ft) in length.[16]

Breeding

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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.^[17]

Diet

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Gopher tortoises are mainlyherbivores that feed on low growing plant life. Their diet consists mostly ofgrasses andlegumes but they will also feed on smallberries andfruits.

Desert tortoise (*Gopherus agassizii*) drinking from roadway inJoshua 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 cm. per year.Gopherus polyphemus live inmesic habitat, where water is available 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 humidmicrohabitats, 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.^[18]^{: 85–87, 94 pp.}

There are many observations ofGopherus 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 byGopherus include supplements for the low levels of calcium, phosphorus, or protein in their diet, and a need forcalcium carbonate in bones, the production of eggshells in females, and growing young. Hypotheses on the consumption of nonfood items such as soil or rocks includemastication orvermifuge for the removal of parasites.^[18]^{: 93 p.}

Conservation

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Texas tortoise (*Gopherus berlandieri*), northernTamaulipas, Mexico

Populations of all six species ofGopherus have declined dramatically. In the past, gopher tortoises were hunted for their meat, which was used instews.^[19] Currently the most significant threat to their survival ishabitat destruction, but the pet trade and collisions with vehicles have also taken their toll.^[4] 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.^[20] 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.^[21] In 2018, the IUCN Tortoise and Freshwater Turtle Specialist Group recommended a re-assessment and re-classification of all sixGopherus species^[22] This reclassification would moveG. 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) andG. polyphemus from Vulnerable (VU) to Endangered (EN).^[22]

Gallery

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A juvenile bolson tortoise (Gopherus flavomarginatus), from Durango, Mexico (7 July 2006)
Desert tortoise (Gopherus agassizii)San Bernardino County, California, US.
Gopher tortoise (Gopherus polyphemus) from Lee County, Florida (18 June 2018)
Gopher tortoise (Gopherus polyphemus) from Volusia County, Florida (14 September 2008)

References

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^Le, M.; Raxworthy, C. J.; McCord, W. P.; Mertz, L. (2006-05-05). "A molecular phylogeny of tortoises (Testudines: Testudinidae) based on mitochondrial and nuclear genes".Molecular Phylogenetics and Evolution.40 (2):517–531.doi:10.1016/j.ympev.2006.03.003.PMID 16678445.
^"About The Gopher tortoise". Gopher Tortoise Council. Archived fromthe original on 2009-09-01. Retrieved2009-07-03.
^Fergus, Charles (2007).Turtles. Stackpole Books. pp. 22–24.ISBN 978-0-8117-3420-2.
^^a ^bFranklin, Carl J. (2007).Turtles: An Extraordinary Natural History 245 Million Years in the Making. MBI Publishing Company. pp. 129–131.ISBN 978-0-7603-2981-8.
^"Fossilworks:Gopherus".fossilworks.org. Retrieved2020-02-29.
^^a ^bUS Geological Survey
^Murphy, Robert; Berry, Kristin; Edwards, Taylor; Leviton, Alan; Lathrop, Amy; Riedle, J. Daren (2011-06-28)."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".ZooKeys (113):39–71.Bibcode:2011ZooK..113...39M.doi:10.3897/zookeys.113.1353.ISSN 1313-2970.PMC 3187627.PMID 21976992.
^^a ^b ^cEdwards T, Karl AE, Vaughn M, Rosen PC, Torres CM, Murphy RW (2016)."The desert tortoise trichotomy: Mexico hosts a third, new sister-species of tortoise in theGopherus morafkai–G. agassizii group".ZooKeys (562):131–158.Bibcode:2016ZooK..562..131E.doi:10.3897/zookeys.562.6124.PMC 4768471.PMID 27006625.
^Gopherus, The Reptile Database
^^a ^bBramble, Dennis M. and J. Howard Hutchison. 2014.Morphology, Taxonomy, and Distribution of North American Tortoises: An Evolutionary Perspective. pages 1-12In, 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
^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
^Reynoso, Víctor-Hugo; Montellano-Ballesteros, Marisol (2004-01-01)."A New Giant Turtle of the GenusGopherus (Chelonia: Testudinidae) from the Pleistocene of Tamaulipas, México, and a Review of the Phylogeny and Biogeography of Gopher Tortoises"(PDF).Journal of Vertebrate Paleontology.24 (4):822–837.doi:10.1671/0272-4634(2004)024[0822:angtot]2.0.co;2.JSTOR 4524778.S2CID 84397134.
^Rostal, David C.; McCoy, Earl D.; Mushinsky, Henry R. (2014-06-13).Biology and Conservation of North American Tortoises. JHU Press.ISBN 9781421413778.
^"Ecological history and latent conservation potential: large and giant tortoises as a model for taxon substitutions"(PDF).
^Woodburne & Golz, 1972
^Esker, Donald A.; Forman, Steve L.; Butler, Dava K. (May 2019)."Reconstructing the mass and thermal ecology of North American Pleistocene tortoises".Paleobiology.45 (2):363–377.Bibcode:2019Pbio...45..363E.doi:10.1017/pab.2019.6.ISSN 0094-8373.S2CID 155216574.
^Landers, L. J.; Garner, J. A. & McRae, W. A. (1980). "Reproduction on Gopher Tortoises in Southwestern Georgia".Herpetological.36: 353–361.
^^a ^bEsque, 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
^"Gopher Tortoise Stew", in:Recipes from Another Time: Savor the flavor of old St. Augustine and try a couple of these original recipes. Smithsonian magazine, October 2001
^"Fence with an "angle" protects gopher tortoises". US Department of Transportation. 2003-04-25. Archived fromthe original on June 25, 2003. Retrieved2010-04-18.
^"Department of the Interior: Fish and Wildlife Service: 50 CFR Part 17"(PDF).Federal Register. Vol. 74, no. 215. 2009-11-09. Retrieved2009-11-13.
^^a ^bRhodin, Anders G. J.; Stanford, Craig B.; Dijk, Peter Paul Van; Eisemberg, Carla; Luiselli, Luca; Mittermeier, Russell A.; Hudson, Rick; Horne, Brian D.; Goode, Eric V.; Kuchling, Gerald; Walde, Andrew (December 2018)."Global Conservation Status of Turtles and Tortoises (Order Testudines)".Chelonian Conservation and Biology.17 (2):135–161.doi:10.2744/CCB-1348.1.ISSN 1071-8443.S2CID 91937716.

Bibliography

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Woodburne, M. O.; Golz, D. J. (1972).Stratigraphy of the Punchbowl Formation, Cajon Valley, southern California.University of California Publications in Geological Sciences. Vol. 92.

External links

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Wikimedia Commons has media related toGopherus.

Wikispecies has information related toGopherus.

Tortoise family

Kingdom:Animalia
Phylum:Chordata
Class:Reptilia
Subclass:Anapsida
Order:Testudines
Suborder:Cryptodira
Superfamily:Testudinoidea
Family:Testudinidae

Genera

Species of thetortoise family

Agrionemys

Russian tortoise

Angulate tortoise

†Hadrianus

†Hesperotestudo percrassa

Pancake tortoise

Leopard tortoise

†Stylemys

Testudo

Phylogenetic arrangement of turtles based onturtles of the world 2017 update: Annotated checklist and atlas of taxonomy, synonymy, distribution, and conservation status. Key: †=extinct.

Testudines

Kingdom:Animalia
Phylum:Chordata
Class:Reptilia
Clade:Diapsida
Order:Testudines

Genus

Chelonioidea
(Sea turtles)

Cheloniidae	†Allopleuron Caretta †Carolinochelys Chelonia †Eochelone Eretmochelys †Gigantatypus †Glarichelys †Itilochelys Lepidochelys †Mexichelys †Miocaretta Natator †Pacifichelys †Syllomus †Tasbacka
Dermochelyidae	†Arabemys †Corsochelys †Cosmochelys Dermochelys †Eosphargis †Mesodermochelys †Psephophorus
	†Euclastes †Peritresius †Procolpochelys †Protosphargis †Puppigerus

Kinosternoidea

Dermatemydidae	Dermatemys
Kinosternidae	Claudius †Hoplochelys Kinosternon Staurotypus Sternotherus

Testudinoidea

Emydidae	†Acherontemys Chrysemys Clemmys Deirochelys Emys Actinemys Emydoidea Glyptemys Graptemys Malaclemys Pseudemys Terrapene Trachemys †Wilburemys
Geoemydidae	Batagur †Banhxeochelys Cuora Cyclemys Geoclemys Geoemyda Hardella Heosemys Leucocephalon Malayemys Mauremys Melanochelys Morenia Notochelys Orlitia Pangshura Rhinoclemmys Sacalia Siebenrockiella Vijayachelys
Platysternidae	Platysternon
Testudinidae	Aldabrachelys Astrochelys Centrochelys Chelonoidis Chersina Cylindraspis †Cymatholcus †Floridemys Geochelone Gopherus †Hadrianus †Hesperotestudo Homopus Indotestudo Kinixys Malacochersus Manouria †Megalochelys †Oligopherus Psammobates Pyxis †Solitudo Stigmochelys †Stylemys Testudo

Trionychia

Carettochelyidae	†Allaeochelys †Anosteira Carettochelys
Trionychidae	Amyda Apalone †Axestemys Chitra Cyclanorbis Cycloderma †Drazinderetes Dogania †Gilmoremys †Hutchemys †Khunnuchelys Lissemys Nilssonia Palea †Palaeoamyda Pelochelys Pelodiscus Rafetus Trionyx
	†Basilochelys †Sinaspideretes

Chelydridae	Chelydra †Chelydrops †Chelydropsis †Emarginachelys †Macrocephalochelys Macrochelys †Planiplastron †Protochelydra
†Nanhsiungchelyidae	†Anomalochelys †Basilemys †Jiangxichelys
†Protostegidae	†Alienochelys †Archelon †Atlantochelys †Bouliachelys †Calcarichelys †Cratochelone †Desmatochelys †Iserosaurus †Notochelone †Ocepechelon †Pneumatoarthrus †Protostega †Rhinochelys †Santanachelys †Terlinguachelys
	†Adocus †Argillochelys †Bashuchelys †Ctenochelys †Prionochelys †Toxochelys

Pleurodira

†Araripemydidae	†Araripemys
†Bothremydidae	†Araiochelys †Arenila †Azabbaremys †Bothremys †Cearachelys †Chedighaii †Chupacabrachelys †Eotaphrosphys †Foxemys †Galianemys †Ilatardia †Inaechelys †Itapecuruemys †Jainemys †Kinkonychelys †Kurmademys †Labrostochelys †Nigeremys †Phosphatochelys †Polysternon †Puentemys †Rosasia †Rhothonemys †Sankuchemys †Taphrosphys †Ummulisani †Zolhafah
Chelidae	Acanthochelys Chelodina Chelus Elseya Elusor Emydura Hydromedusa †Lomalatachelys Mesoclemmys Myuchelys Phrynops Platemys †Prochelidella Pseudemydura Rheodytes Rhinemys †Yaminuechelys
Pelomedusidae	Pelomedusa Pelusios
Podocnemididae	†Albertwoodemys †Bauruemys †Brontochelys †Caninemys †Carbonemys †Cerrejonemys †Cordichelys Erymnochelys †Lapparentemys †Latentemys Peltocephalus Podocnemis †Stupendemys
†Sahonachelyidae	†Sahonachelys †Sokatra

	†Caribemys †Caririemys †Tacuarembemys

Phylogenetic arrangement of turtles based onTurtles of the World 2017 Update: Annotated Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation Status. † = extinct.

See alsoList of Testudines families

Taxon identifiers
Gopherus	Wikidata:Q1012045 Wikispecies:Gopherus BOLD:191000 CoL:634MS EoL:58352 GBIF:2441807 iNaturalist:40082 ITIS:173855 NCBI:38771 Open Tree of Life:831058 Paleobiology Database:37741

Authority control databases
National	Israel
Other	Yale LUX

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