gopherus is a genus of fossorial tortoises 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 asian genus manouria .

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 their ability to dig large , deep burrows ; those of the gopher tortoise can be up to 40 feet ( 12 m ) in length and 10 feet ( 3.0 m ) in depth .

these burrows are used by a variety of other species , including mammals , other reptiles , amphibians , and birds .

gopher tortoises are 20 – 50 cm ( 7.9 – 19.7 in ) in length , depending on the species .

all five species are found in xeric habitats .

gopherus

gopherus agassizii can hybridize with gopherus berlandieri ( edwards et al . 2010 ) .
s1 fig . divergence summary of repetitive elements in the genome of gopherus agassizii .
conservative estimate of the distribution of gopherus evgoodei in mexico indicated by diagonal lines . desert tortoise range limit , modified based on our field sampling , from . squares indicate museum and literature records of occurrence of gopherus spp . circles are sample locations from for both gopherus morafkai ( black ) and gopherus evgoodei ( white ) . localities in the sinaloan thornscrub - sonoran desertscrub ecotone indicated by split circles , which indicate the occurrence of both gopherus evgoodei and gopherus morafkai genotypes and / or hybrids .
detailed view of the anal scutes of the holotype of gopherus morafkai , cas 33867 .
contributions of private landowners to the conservation of the gopher tortoise ( gopherus polyphemus ) .
parentage assignment among breeding adult bolson tortoises ( gopherus flavomarginatus ) for yearly cohorts of hatchlings .
s6 table . enriched go categories using unique , fixed variants per taxon for gopherus agassizii .
s7 table . enriched go categories using unique , fixed variants per taxon for gopherus morafkai .
s8 table . enriched go categories using unique , fixed variants per taxon for gopherus evgoodei .
ventral surface of the right rear foot of the holotype of gopherus evgoodei , amnh r64160 .
right , lateral view of the head of the holotype of gopherus morafkai , cas 33867 .
gopherus agassizii ( cooper 1861 ) ( partim ) . generic reassignment by stejneger ( 1893 )
longevity and growth strategies of the desert tortoise ( gopherus agassizii ) in two american deserts .
what ' s in a name ? gopherus by mary cohen & michael j . connor - cttc
dorsal view of the lectotype of gopherus agassizii , usnm 7888 . black bar is 3 cm .
ventral view of the holotype of gopherus agassizii , usnm 7888 . black bar is 3 cm .
anterior view of the holotype of gopherus agassizii , usnm 7888 . black bar is 3 cm .
implications of anthropogenic landscape change on inter - population movements of the desert tortoise ( gopherus agassizii ) .
mojave population of the desert tortoise ( gopherus agassizii ) . 5 - year review : summary and evaluation
regional - scale estimation of density and habitat use of the desert tortoise ( gopherus agassizii ) in arizona .
genetic and morphological assessment of an unusual tortoise ( gopherus agassizii ) population in the black mountains of arizona .
contributions of private landowners to the conservation of the gopher tortoise ( gopherus polyphemus ) . - pubmed - ncbi
dorsal view of the holotype of gopherus evgoodei , amnh r64160 . scale bar 50 mm in 10 mm increments .
ventral view of the holotype of gopherus evgoodei , amnh r64160 . scale bar 50 mm in 10 mm increments .
habitat use by desert tortoises ( gopherus agassizii ) on alluvial fans in the sonoran desert , south - central arizona .
the desert tortoise trichotomy : mexico hosts a third , new sister - species of tortoise in the gopherus morafkai–g . agassizii group
s3 dataset . assigned 1 : 1 orthologs between agassiz’s desert tortoise ( gopherus agassizii ) and chicken ( gallus gallus ) .
s5 dataset . assigned 1 : 1 orthologs between agassiz’s desert tortoise ( gopherus agassizii ) and human ( homo sapiens ) .
physical characteristics and patterns of utilization of cover sites used by gopherus agassizi in southern nevada . desert tortoise council , proc . symp
class : reptilia order : chelonia suborder : cryptodira super family : testudinoidea family : testudinidae genus : gopherus species : g . agassizii
s4 dataset . assigned 1 : 1 orthologs between agassiz’s desert tortoise ( gopherus agassizii ) and green anole ( anolis carolinensis ) .
mertens and wermuth ( 1955 ) and wermuth and mertens ( 1961 ) were unimpressed by the extent of morphological differentiation among north american gopherus and impressed by the reports of hybrids . while recognizing long - term isolation , they recognized only one species , gopherus polyphemus , stating “da sich die einzelnen formen der gopherschildkröten äußberlich nur wenig unterscheiden , deutlich geographisch vikariieren und mehreren veröffentlichungen zufolge auch zu verbastardieren scheinen , sind sie hier als unterarten aufgeführt” ( wermuth and mertens 1961 : 172 ) . in doing so , they considered gopherus agassizii to be a subspecies of gopherus polyphemus , gopherus polyphemus agassizii ( cooper ) . their taxonomic arrangement was rarely , if ever , followed .
gopherus evgoodei edwards , karl , vaughn , rosen , meléndez - torres & murphy 2016 xerobates agassizii cooper 1861 ( partim ) gopherus agassizii — stejneger 1893 ( partim ) scaptochelys agassizii — bramble 1982 ( partim ) xerobates lepidocephalus — ottley & velázques - solis 1989 ( in error ) xerobates lepidocephalus — crumly & grismer 1994 ( in error ) gopherus morafkai — murphy et al . 2011 ( partim )
s2 dataset . assigned 1 : 1 orthologs between agassiz’s desert tortoise ( gopherus agassizii ) and chinese softshell turtle ( pelodiscus sinensis ) .
to evaluate the validity of gopherus lepidocephalus and to confirm the geographic origin of gopherus agassizii , we obtained mitochondrial dna sequences from both type specimens . this kind of analysis could not detect hybrids because the mitochondrial genome is inherited only maternally . however , if gopherus lepidocephalus has its origin in the mojave desert , then the name will persist as a junior synonym of gopherus agassizii regardless of whether it is a hybrid or not . alternatively , if the maternal lineage is from a sonoran desert tortoise , then the possible hybrid state would create another problem to be solved . finally , if the lineage was new and divergent , then perhaps gopherus lepidocephalus was native to the peninsula .
mushinsky , h . , d . wilson , e . mccoy . 1994 . growth and sexual dimorphism of gopherus polyphemus in central florida .
s1 dataset . assigned 1 : 1 orthologs between agassiz’s desert tortoise ( gopherus agassizii ) and western painted turtle ( chrysemys picta bellii ) .
the desert tortoise trichotomy : mexico hosts a third , new sister - species of tortoise in the gopherus morafkai - g . agassizii group .
distribution : see map in murphy et al . 2016 : 133 . hybridization : gopherus evgoodei hybridizes with g . morafkai in c sonora .
in terms of generic allocation , crumly ( 1994 ) aptly notes that the genus xerobates cannot be diagnosed morphologically owing to intraspecific variation . thus , he refers xerobates agassizii back to gopherus agassizii ( cooper , 1861 ) . symplesiomorphies are used by bramble ( 1982 ) to define scaptochelys , a practice that contravenes the principles of phylogenetic systematics . although morphological evidence does not unite gopherus agassizii and gopherus berlandieri , molecular evidence does ( lamb and lydeard 1994 ) . and although it is possible to recognize xerobates for the extant species gopherus agassizii and gopherus berlandieri , the phylogenetic relationships among extinct species ( reynoso and montellano - ballesteros 2004 ) preclude monophyly of the two genera . thus , xerobates should not be recognized .
wirt , b . 2011 . desert tortoise - gopherus agassizii . sonoran herpetologist 24 ( 6 ) : 56 - 58 . - get paper here
stitt , e . , c . schwalbe , d . swann . 2004 . gopherus agassizii ( desert tortoise ) . association with africanized bees .
gene diversity ( a ) and allelic richness ( b ) per locus and population of bolson tortoises ( gopherus flavomarginatus ) for 11 str loci .
van dijk pp , flores - villela o ( 2007 ) gopherus flavomarginatus iucn 2013 : iucn red list of threatened species . urltoken : iucn .
edwards , t . , e . stitt , c . schwalbe , d . swann . 2004 . gopherus agassizii ( desert tortoise ) . movement .
the genus name gopherus was coined by the naturalist c . s . rafinesque in 1816 but first published in 1832 to describe the north american tortoises . gopherus is derived from the word gopher , which was already in common use in the us to describe burrowing animals such as the gopher tortoise at the time .
woodbury , angus munn ; hardy , ross 1948 . studies of the desert tortoise , gopherus agassizii . ecological monographs 18 ( 2 ) : 146 - 200
2013 .
gopher tortoise : gopherus polyphemus
( on - line ) . florida fish and wildlife conservation commission . accessed november 29 , 2013 at urltoken .
the desert tortoise trichotomy : mexico hosts a third , new sister - species of tortoise in the gopherus morafkai - g . agassizii group . - pubmed - ncbi
egg size and annual egg production by female desert tortoises ( gopherus agassizii ) : the importance of food abundance , body size , and date of egg shelling .
auffenberg , w . , and r . franz . 1978 . gopherus flavomarginatus . catalogue of american amphibians and reptiles 214 : 1 - 2 . - get paper here
miller , loye 1955 . further observations on the desert tortoise , gopherus agassizi , of california . copeia 1955 ( 2 ) : 113 - 118 - get paper here
sullivan , brian k . and elizabeth sulivan . 2014 . gopherus morafkai ( sonoran desert tortoise ) drinking behavior . herpetological review 45 ( 3 ) : 483 - 484
sullivan , brian k . and elizabeth sullivan . 2016 . gopherus morafkai ( sonoran desert tortoise ) drinking behavior . herpetological review 47 ( 1 ) : 123 - 124
s4 fig . representation of gene ontology terms for biological processes shared by genes with ka / ks > 1 in the pairwise comparison of gopherus agassizii and gallus gallus .
desert tortoises ( testudines ; testudinidae ; gopherus agassizii group ) have an extensive distribution throughout the mojave , colorado , and sonoran desert regions . not surprisingly , they exhibit a tremendous amount of ecological , behavioral , morphological and genetic variation . gopherus agassizii was considered a single species for almost 150 years but recently the species was split into the nominate form and morafka ' s desert tortoise , gopherus morafkai , the latter occurring south and east of the colorado river . whereas a large body of literature focuses on tortoises in the united states , a dearth of investigations exists for mexican animals . notwithstanding , mexican populations of desert tortoises in the southern part of the range of gopherus morafkai are distinct , particularly where the tortoises occur in tropical thornscrub and tropical deciduous forest . recent studies have shed light on the ecology , morphology and genetics of these southern ' desert ' tortoises . all evidence warrants recognition of this clade as a distinctive taxon and herein we describe it as gopherus evgoodei sp . n . the description of the new species significantly reduces and limits the distribution of gopherus morafkai to desertscrub habitat only . by contrast , gopherus evgoodei sp . n . occurs in thornscrub and tropical deciduous forests only and this leaves it with the smallest range of the three sister species . we present conservation implications for the newly described gopherus evgoodei , which already faces impending threats .
our investigation of the taxonomy of agassiz’s land tortoise resolved many issues . the publication date has been given in error as 1863 since its first citation . the type series was likely collected by cooper from near soda lake , california , and not elsewhere . only one of the three original cotypes exists , usnm 7888 , and it was designated as the lectotype . our mtdna sequence data from the lectotype confirmed that it was from california , not arizona . further , mtdna sequence data from the holotype of gopherus lepidocephalus placed its origin to the mojavian population , rather than the sonoran desert of either arizona or mexico . genetic , morphological and ecological data confirmed the existence of at least two species contained within gopherus agassizii . the sonoran population is named as a new species , gopherus morafkai , morafka ` s desert tortoise . the recognition of gopherus morafkai reduces the range of gopherus agassizii to occupying about 30 % of its former range . given drastic population declines in gopherus agassizii during the past few decades , it might be endangered .
2012 .
anage entry for gopherus polyphemus
( on - line ) . anage : the animal ageing and longevity database . accessed december 01 , 2013 at urltoken .
to cite this page : lazzari , a . 2017 .
gopherus polyphemus
( on - line ) , animal diversity web . accessed july 09 , 2018 at urltoken
auffenberg w ; franz r 1978 . gopherus agassizii ( cooper ) . desert tortoise . catalogue of american amphibians and reptiles ( 212 : 1 - 2 - get paper here
to cite this page : crozier , g . 1999 .
gopherus agassizii
( on - line ) , animal diversity web . accessed july 09 , 2018 at urltoken
diversity indices for 11 microsatellite loci in 3 sample populations of bolson tortoises , gopherus flavomarginatus ; appleton ranch , durango mexico , and hybrid population at the el paso zoo .
innes , r . 2009 .
gopherus polyphemus
( on - line ) . usda forest service : fire effects information system . accessed december 02 , 2013 at urltoken .
winchell , s . 2006 . tortoises of north america - an overview of the genus gopherus . reptilia ( gb ) ( 49 ) : 9 - 15 - get paper here
bruekers , jaco 2004 . in der sporen van de amerikaanse woestijnschildpad , gopherus agassizii ( cooper 1863 ) . lacerta 62 ( 5 ) : 198 - 2005 - get paper here
crumly , c . r . 1994 . phylogenetic systematics of north american tortoises ( genus gopherus ) : evidence for their classification . fish and wildlife research 13 : 7 - 32
dna was extracted a minimum of three times for the lectotype of gopherus agassizii and once for the holotype of gopherus lepidocephalus . to avoid any possibility of cross contamination , final extractions were done in isolation of one another . amplification and sequencing were also done independently for both strands . desert tortoise sequences were confirmed using a blast search of the ncbi database .
phylogenetic systematics of north american tortoises ( genus gopherus ) : evidence for their classification . in : bury rb , germano dj ( eds ) , biology of north american tortoises .
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 .
gonzález trápaga , r . & aguirre léon , g . 2006 . gopherus flavomarginatus - the bolson tortoise . reptilia ( gb ) ( 49 ) : 22 - 27 - get paper here
peterson , charles c . 1994 . different rates and causes of high mortality in two populations of the threatened desert tortoise gopherus agassizii . biological conservation 70 ( 2 ) : 101 - 108
averill - murray , r . , a . averill - murray . 2005 . regional - scale estimation of density and habitat use of the desert tortoise ( gopherus agassizii ) in arizona .
nomenclatural stability for gopherus agassizii was maintained until bramble ( 1982 ) revised the genus using both extant and extinct species . he discovered two groups and erected the genus scaptochelys for gopherus agassizii and gopherus berlandieri . the type species was designated as xerobates agassizii cooper , 1863 [ sic ] . thus , gopherus agassizii was referred to as scaptochelys agassizii . shortly thereafter , bour and dubois ( 1984 ) reported that scaptochelys was a junior synonym of genus xerobates agassiz , 1857 , whose type species was xerobates berlandieri agassiz , 1857 , by subsequent designation ( brown 1908 ) . because bramble ( 1982 ) resolved scaptochelys agassizii as the sister group of xerobates berlandieri , bour and dubois ( 1984 ) referred scaptochelys agassizii back to xerobates agassizii cooper , 1863 [ sic ] .
female gopherus morafkai mature at larger sizes ( 220 mm carapace length ) ( averill - murray 2002 ) than does gopherus agassizii ( 176–190 mm carapace length ) ( turner et al . 1986 ; germano 1994a ; karl 1998 ) . clutch sizes between the two species are similar ( averill - murray 2002 ) , but gopherus morafkai only produces 1 clutch every 1–2 yr ( averill - murray 2002 ) while gopherus agassizii may produce 1–3 clutches every year ( turner et al . 1986 ; wallis et al . 1999 ) . harsher , more arid climates in the mojave desert may have led to increased female reproductive investment to offset hatchling and juvenile mortality ( heppell 1998 ; hellgren et al . 2000 ) , but information is limited for juvenile tortoises of both species .
burke , r . , m . ewert , j . mclemore , d . jackson . 1996 . temperature - dependent sex determination and hatching success in the gopher tortoise ( gopherus polyphemus ) .
gonzález trápaga , r . & aguirre león , g . 2006 . gopherus flavomarginatus , die mexikanische gopherschildkröte . reptilia ( münster ) 11 ( 62 ) : 28 - 34 - get paper here
spencely , ashley , jeremy mack and kristin h . berry . 2015 . gopherus agassizii ( agassiz ' s desert tortoise ) attempted predation . herpetological review 46 ( 3 ) : 422 - 423
( a ) estimated divergence of gopherus agassizii and chrysemys picta bellii without constraint . ( b ) fixed age of node 9 ( see s2 fig . ) at 74 . 2 million years .
legler , john m . ; webb , robert g . 1961 . remarks on a collection of bolson tortoises , gopherus flavomarginatus . herpetologica 17 ( 1 ) : 26 - 37 - get paper here
morafka , d . j . 1982 . the status and distribution of the bolson tortoise ( gopherus flavomarginatus ) . u . s . fish and wildlife service research report 12 : 71 - 94 .
berry , kristin h . , shields , tim and jacobson , elliott r . 2016 . gopherus agassizii ( mohave desert tortoise ) probable rattlesnake envenomation herpetological review 47 ( 4 ) : 652 - 653
martin , b . 1996 . the desert tortoise ( gopherus agassizii ) in a desert grassland community in southern arizona . sonoran herpetologist 9 ( 4 ) : 31 - 35 . - get paper here
the generic allocation of the agassiz land - tortoise , the desert tortoise , has occasionally changed . cope ( 1875 ) transferred xerobates agassizii to the genus testudo , as testudo agassizii , in his checklist of north american amphibians and reptiles but without comment or justification . presumably , this determination followed the generic allocation of gray ( 1870 ) and certainly this was not an oversight as cope ( 1880 ) repeated the generic allocation for gopherus berlandieri . the next taxonomic change was made by stejneger ( 1893 ) in his discussion of the fauna of death valley . he considered the californian tortoise to be distinct from gopherus berlandieri and to belong to the north american genus gopherus rafinesque 1832 , as “ gopherus agassizii ( cooper ) ” ( stejneger 1893 : 160 ) . this generic allocation was stable for almost 100 years .
winchell , s . & love , b . 2006 . die landschildkröten nordamerikas . ein überblick über die gattung gopherus . reptilia ( münster ) 11 ( 62 ) : 20 - 27 - get paper here
lovich , jeffrey e . 2011 . gopherus agassizii ( desert tortoise ) and crotalus ruber ( red diamond rattlesnake ) burrow co - occupancy . herpetological review 42 ( 3 ) : 421 - get paper here
walde , andrew d . , angela m . walde and a . peter woodman . 2016 . gopherus agassizii ( mojave desert tortoise ) burrow associate . herpetological review 47 ( 1 ) : 121 - 122
vaughn sl ( 1984 ) home range and habitat use of the desert tortoise ( gopherus agassizii ) in the picacho mountains , pinal county , arizona . m . sc . thesis , arizona state university .
martin be ( 1995 ) ecology of the desert tortoise ( gopherus agassizii ) in a desert - grassland community in southern arizona . m . sc . thesis , university of arizona , tucson , az .
to determine gopherus evgoodei was in fact a new species , edwards sampled 233 wild desert tortoises of both the sonoran and sinaloan lineage and performed dna analysis on these reptiles and compared dna with that of the gopherus evgoodei and found distinct differences in the dna . edwards said that the tortoise species changes with the vegetation . those to the north are the sonoran desert tortoises while the south holds the thornscrub tortoise .
zylstra , e . 2007 . the bolson tortoise ( gopherus flavomarginatus ) : king of the ( re - ) wild frontier . sonoran herpetologist 20 ( 5 ) : 50 - 54 . - get paper here
(
anage entry for gopherus polyphemus
, 2012 ;
gopher tortoise management plan
, 2012 ; conant and collins , 1998 ; ernst and lovich , 2009 ; mushinsky , et al . , 1994 )
moll , e . o . 2004 . gopherus agassizi ( cooper , 1863 ) – desert tortoise . patronyms of the pioneer west . sonoran herpetologist 17 ( 5 ) : 50 - 53 . - get paper here
smith , amanda l . , laura a . tennant and terence r . arundal . 2015 . gopherus agassizii ( agassiz ' s desert tortoise ) mechanical injury . herpetological review 46 ( 3 ) : 423 - 424
henderson , r . a . , s . r . puffer and j . e . lovich 2016 . gopherus agassizii ( mohave desert tortoise ) nest depredation . herpetological review 47 ( 3 ) : 446 - 447 .
walde , andrew d . , meagan l . harless , david k . delaney and larry l . pater . 2006 . gopherus agassizii diet . herpetological review 37 ( 1 ) : 77 - 78 - get paper here
walde , angela m . , andrew d . walde and charlie jones . 2014 . gopherus agassizii ( mohave desert tortoise ) and crotalus mitchelli ( speckled rattlesnake ) burrow associate . herpetological review 45 ( 4 ) : 688
webster , timothy h . ; greer a . dolby , melissa wilson sayres , kenro kusumi 2017 . improved draft of the mojave desert tortoise genome , gopherus agassizii , version 1 . 1 peerj preprints - get paper here
several observations suggested the absence of dna contamination . first , amplification of dna from the two type specimens resulted in differing fragment lengths . primers used for the lectotype of gopherus agassizii did not amplify dna from the holotype of gopherus lepidocephalus . thus , it is exceptionally unlikely that contamination occurred between these two species . neither type specimen had dna extracted along with other samples of gopherus ; all comparative samples were downloaded from genbank . thus , cross - contamination outside of this project was not possible . finally , dna extracted in isolation of the other type precluded the possibility of contamination . consequently , all evidence suggested that the sequence data were obtained from the respective specimens .
berry et al . ( 2002 ) summarize data suggesting that the desert tortoise , gopherus agassizii ( cooper ) , of the southern united states and northwestern mainland mexico is a composite of at least two and possibly four species . they note that much work remains to be accomplished before formally recognizing any new species . this task is more complex than originally imagined , in part because of a convoluted taxonomy plagued with uncertainties and problems . our reviews of several conundrums obtain the background data required to untangle a knot of confusion and make some decisions and recommendations . the greatest problem concerns the identities of true gopherus agassizii and the enigmatic gopherus lepidocephalus ( ottley et velázques solis ) .
legler , j . m . 1959 . a new tortoise , genus gopherus , from north - central mexico . univ . kansas publ . mus . nat . hist . 11 : 335 - 343 . - get paper here
walde , andrew d . ; andrea currylow 2015 . gopherus agassizii ( mojave desert tortoise ) and coleonyx variegatus variegatus ( desert banded gecko ) . spring burrow cohabitation . herpetology notes 8 : 501 - 502 - get paper here
sullivan , b . k . , and e . a . sullivan . 2015 . winter basking by hatchling sonoran desert tortoises , gopherus morafkai . sonoran herpetologist 28 ( 1 ) : 10 - 11 . - get paper here
currylow , andrea , andrew d . walde and angela m . walde . 2016 . gopherus agassizii ( mojave desert tortoise ) and coluber flagellum piceus ( red racer ) burrow associates . herpetological review 47 ( 1 ) : 122 - 123
curtin , a . j . ; g . r . zug ; j . r . spotila 2009 . longevity and growth strategies of the desert tortoise ( gopherus agassizii ) in two american deserts . journal of arid environments 73 : 463–471
the questioned identity of gopherus lepidocephalus has now been sufficiently answered to address its taxonomic status . the name is a junior synonym of gopherus agassizii . whether the holotype is a hybrid or not is taxonomically irrelevant because the maternal lineage had an origin in the mojave desert population . nevertheless , three questions remain . first , is the holotype of gopherus lepidocephalus a hybrid individual ? this could explain its uniqueness ( ottley and velázques solis 1989 ) as well as it association with sonora , mexico ( crumly and grismer 1994 ) . second , the question remains as to whether gopherus morafkai consists of two forms that warrant recognition at the species level : morafka’s desert tortoise and a potentially new sinaloan thornscrub tortoise ( lamb et al . 1989 ) . currently , we are examining the spatial overlap of several genotypes at the eastern and southern boundaries of sonoran desert scrub in sonora , mexico to better understand the evolutionary drivers responsible for shaping the genetic diversity of gopherus morafkai , and to evaluate the possibility that the species is a composite of two cryptic species . finally , it is critical to evaluate ontogenetic development in both species . this may vary geographically within species as well as with nutrition and other environmental parameters .
we investigate a cornucopia of problems associated with the identity of the desert tortoise , gopherus agassizii ( cooper ) . the date of publication is found to be 1861 , rather than 1863 . only one of the three original cotypes exists , and it is designated as the lectotype of the species . another cotype is found to have been destroyed in the 1906 san francisco earthquake and subsequent fire . the third is lost . the lectotype is genetically confirmed to be from california , and not arizona , usa as sometimes reported . maternally , the holotype of gopherus lepidocephalus ( ottley & velázques solis . 1989 ) from the cape region of baja california sur , mexico is also from the mojavian population of the desert tortoise , and not from tiburon island , sonora , mexico as previously proposed . a suite of characters serve to diagnose tortoises west and north of the colorado river , the mojavian population , from those east and south of the river in arizona , usa , and sonora and sinaloa , mexico , the sonoran population . species recognition is warranted and because gopherus lepidocephalus is from the mojavian population , no names are available for the sonoran species . thus , a new species , gopherus morafkai sp . n . , is named and this action reduces the distribution of gopherus agassizii to only 30 % of its former range . this reduction has important implications for the conservation and protection of gopherus agassizii , which may deserve a higher level of protection .
auffenberg , w . , & franz , r . 1978 . gopherus rafinesque : gopher tortoises . catalogue of american amphibians and reptiles ( 211 ) : 1 - 2 [ species accounts in nos . 212 - 215 ] - get paper here
ureña - aranda , cinthya alejandra ; de los monteros , alejandro espinosa 2012 . the genetic crisis of the mexican bolson tortoise ( gopherus flavomarginatus : testudinidae ) . amphibia - reptilia 33 ( 1 ) : 45 - 53 - get paper here
the new species is a patronym for the late professor david joseph morafka in recognition of his many contributions to the biology and conservation of the species of gopherus and his unsurpassed way of facilitating research , even among researchers with very different perspectives .
gray , k . m . , and r . steidl . 2014 . effects of buffelgrass invasion on density and condition of sonoran desert tortoises ( gopherus morafkai ) . sonoran herpetologist 27 ( 4 ) : 91 - 95 . - get paper here
arizona - sonora desert museum ' s tortoise adoption program , 2003 .
the desert tortoise ( gopherus agassizii ) : a natural history
( on - line ) . arizona - sonora desert museum . accessed september 27 , 2005 at urltoken .
( a ) commands used for the various steps in the genome assembly process . ( b ) commands used to filter the variant file ( . vcf ) from freebayes using snpsft , and to intersect it with the gopherus agassizii annotation using bedtools .
us fish and wildlife service . range - wide monitoring of the mojave desert tortoise ( gopherus agassizii ) : 2013 and 2014 annual reports . report by the desert tortoise recovery office , us fish and wildlife service , reno , nevada . 2015 .
common names do not enjoy precedence and they can create much unnecessary confusion . historically , the species of gopherus were commonly referred to simply as gophers , a word that normally refers to mammals . now that gopherus morafkai is recognized , the desert tortoise requires two common names . gopherus agassizii could be referred to as the mojavian desert tortoise , yet this is inaccurate because the species also occurs within the sonoran desert of california . therefore , we prefer to call it agassiz´s desert tortoise . this name also serves to retain the original designation of cooper ( 1861 ) . similarly , gopherus morafkai occurs in the mojave desert of arizona , the sonoran desert of arizona , usa and sonora , mexico and in sinaloan thornscrub , but not in the mojave and sonoran deserts of california . therefore , the term sonoran desert tortoise is inaccurate . consequently , we prefer to call this species morafka´s desert tortoise . these common names will serve to exclude the species from other desert tortoises in the genus testudo .
however , when the texas and desert tortoises were first described in the nineteenth century they had been given the genus name xerobates ( from the greek xero meaning
dry
and bates meaning
one who walks or haunts
) . under the rules used in assigning scientific names xerobates took priority over scaptochelys because it had been used first . so , the
new
name of the texas tortoise became xerobates berlandieri and the desert tortoise became xerobates agassizii . the florida gopher tortoise and the closely related mexican bolson tortoise remained in the genus gopherus . however , not all authorities accept the new genus ( it took 100 years or so before even gopherus was widely recognized as valid ! ) . this confusion explains why the desert tortoise is sometimes listed as gopherus ( xerobates ) agassizii , a technically
untidy
moniker because it suggests that xerobates is a sub - genus of gopherus and not a separate genus . no doubt this will be clarified in the future when more is known about these tortoises .
hagerty , b . e . ; peacock , m . m . ; kirchoff , v . s . & tracy , c . r . 2008 . polymorphic microsatellite markers for the mojave desert tortoise , gopherus agassizii . molecular ecology resources 8 : 1149–1151
llamas , macario ; christina la croix , elizabeth williams , and william perry baker 2017 . hematological values in captive desert tortoises ( gopherus agassizii ) from maricopa county , arizona . ircf reptiles & amphibians 24 ( 3 ) : 187–190 - get paper here
moll , d . 2008 . dietary characteristics and seed germination influences of desert tortoises ( gopherus agassizii ) in a very wet year ( 1998 ) in northeastern sonoran desert . sonoran herpetologist 21 ( 8 ) : 86 - 89 . - get paper here
babb , randall d . , thomas c . brennan , david d . kandiyeli , christina m . akins and thomas r . jones . 2013 . geographic distribution : gopherus morafkai ( sonoran desert tortoise ) . herpetological review 44 ( 4 ) : 623
distribution of the desert tortoises aligned with gopherus agassizii . the locality of byu 39706 from baja california sur is shown as a black dot . the location of the hybrid population described in mcluckie et al . ( 1999 ) is shown as a star .
the evolutionary species concept ( simpson 1961 ; wiley 1978 ) suggests that the sonoran population of the desert tortoise should be recognized as a new taxon . frost and hillis ( 1990 ) effectively argue that subspecies should not be recognized for continuously distributed species ; we agree . given these two observations , at least two species of desert tortoise should be recognized . the dna sequence data exclude application of the available name gopherus lepidocephalus for the sonoran desert population of gopherus that occurs west and south of the colorado river and they confirm that the lectotype of gopherus agassizii is from the mojave desert , and not arizona . because no names are available for the tortoise population occurring in the sonoran desert south and east of the colorado river , we describe it as a new species .
curtin , amanda j . ; , george r . zug , , philip a . medica , james r . spotila 2008 . assessing age in the desert tortoise gopherus agassizii : testing skeletochronology with individuals of known age . endang species res 5 : 21 - 27
gopherus evgoodei differs from other species of desert tortoises in often having yellow / orange integument ( skin ) and shell . i rancho el divisadero ii–iii rancho las cabras ; and iv–v rancho la sierrita near alamos , sonora , mexico ( in tropical deciduous forest ) .
morafka dj , adest ga , aguirre g , recht m ( 1981 ) the ecology of the bolson tortoise gopherus flavomarginatus . in : barbault r , halffter g , editors . ecology of the chihuahuan desert : mexico df : instituto de ecología . pp . 35–78 .
adest , g . a . ; aguirre , g . ; morafka , d . j . ; jarchow , j . v . 1989 . bolson tortoise ( gopherus flavomarginatus ) conservation : i . life history . vida sylvestre neotropical 2 ( 1 ) : 7 - 13
walde , andrew d . ; andrea currylow , angela m . walde 2015 . discovery of a new burrow associate of the desert tortoise ( gopherus agassizii ) , the long - nosed leopard lizard ( gambelia wislizenii ) herpetology notes 8 : 107 - 109 - get paper here
jamie c . moon , earl d . mccoy , henry r . mushinsky , stephen a . karl ; multiple paternity and breeding system in the gopher tortoise , gopherus polyphemus , journal of heredity , volume 97 , issue 2 , 1 march 2006 , pages 150–157 , urltoken
averill - murray , roy c . and bridgette e . hagerty 2014 . translocation relative to spatial genetic structure of the mojave desert tortoise , gopherus agassizii . chelonian conservation and biology jul 2014 , vol . 13 , no . 1 : 35 - 41 . - get paper here
citation : edwards t , cox ec , buzzard v , wiese c , hillard ls , murphy rw ( 2014 ) genetic assessments and parentage analysis of captive bolson tortoises ( gopherus flavomarginatus ) inform their “rewilding” in new mexico . plos one 9 ( 7 ) : e102787 . urltoken
the flat shell profile / shape of carapace generally distinguishes gopherus evgoodei from other species of desert tortoises . live , wild - caught individuals from ( i–iv ) rancho las cabras and ( v–vi ) rancho la sierrita near alamos , sonora , mexico ( in tropical deciduous forest ) .
gopherus evgoodei differs from other species of desert tortoises in having a very short tail . i rancho el chupadero east of guaymas ( in thornscrub habitat ) ; ii rancho las cabras ; and iii–i rancho la sierrita near alamos , sonora , mexico ( in tropical deciduous forest ) .
tortoises are any of the land - dwelling turtles of the family testudinidae . the desert tortoise is one of four species of the genus gopherus , known collectively as gopher tortoises . gopher tortoises are characterized by brown shells 8 - 15 inches long with flattened front limbs adapted for burrowing .
murphy , r . w . 2011 . 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 . sonoran herpetologist 24 ( 7 ) : 71 - 72 . - get paper here
bridges , andy ; heather l . bateman , audrey k . owens , cristina a . jones and william miller 2016 . microhabitat selection of juvenile sonoran desert tortoises ( gopherus morafkai ) in central arizona . chelonian conservation and biology 15 ( 2 ) : 219 - 230 - get paper here
s8 dataset . output from ka / ks analysis for all genes compared between agassiz’s desert tortoise ( gopherus agassizii ) and western painted turtle ( chrysemys picta bellii ) , chinese softshell turtle ( pelodiscus sinensis ) , green sea turtle ( chelonia mydas ) , and chicken ( gallus gallus ) .
take the desert tortoise , known to many of us as gopherus agassizii . gopherus is a uniquely north american genus consisting of tortoises that are structurally adapted to a burrowing and digging lifestyle . three other species of gopher tortoise are alive today - the texas tortoise ( berlandieri ) , the mexican bolson tortoise ( flavomarginatus ) , and the florida gopher tortoise ( polyphemus ) . over the years , experts in the field came to realize that the genus gopherus includes two sorts of tortoise , separable on the basis of differences in their bone and shell structure . the desert and texas tortoises fall in one group and the gopher and bolson tortoise in the other . in 1982 , john bramble suggested that these differences were great enough to warrant recognition of two separate genera and proposed the genus scaptochelys ( from the greek meaning digger - tortoise ) that would include the desert and texas tortoises .
hazard , lisa c . ; danielle r . shemanski , and kenneth a . nagy 2010 . nutritional quality of natural foods of juvenile and adult desert tortoises ( gopherus agassizii ) : calcium , phosphorus , and magnesium digestibility . journal of herpetology 44 ( 1 ) : 135–147 - get paper here
a desert tortoise ( gopherus agassizii ) diet is comprised mainly of safe grasses and weeds , leafy greens , with small amounts of hard vegetables and moist fruits . a good basic salad can be prepared a week in advance and fed daily with selections from the following served in addition to it .
berry , kristin h . ; lisa m . lyren , julie l . yee , and tracy y . bailey 2014 . protection benefits desert tortoise ( gopherus agassizii ) abundance : the influence of three management strategies on a threatened species . herpetological monographs , 28 : 66 - 92 - get paper here
medica , philip a . ; kenneth e . nussear , todd c . esque , and mary b . saethre 2012 . long - term growth of desert tortoises ( gopherus agassizii ) in a southern nevada population . journal of herpetology 46 ( 2 ) : 213 - 220 . - get paper here
morafka dj ( 1988 ) part iii . historical biogeography of the bolson tortoise . in : morafka dj , mccoy cj , editors . the ecogeography of the mexican bolson tortoise ( gopherus flavomarginatus ) : derivation of its endangered status and recommendations for its conservation : annals of carnegie museum . pp . 47–72 .
s6 dataset . ortholog groups shared between agassiz’s desert tortoise ( gopherus agassizii ) , western painted turtle ( chrysemys picta bellii ) , chinese softshell turtle ( pelodiscus sinensis ) , chicken ( gallus gallus ) , alligator ( alligator mississippiensis ) , green anole ( anolis carolinensis ) and human ( homo sapiens ) .
mcluckie , ann m . ; lamb , trip ; schwalbe , cecil r . ; mccord , robert d . 1999 . genetic and morphometric assessment of an unusual tortoise ( gopherus agassizii ) population in the black mountains of arizona . journal of herpetology 33 ( 1 ) : 36 - 44 - get paper here
gopherus morafkai occurs naturally east and south of the colorado river in arizona , as well as in sonora , including tiburon island , and sinaloa on the west side of the sierra madre occidental , mexico ( berry et al . 2002 ) . the species appears to have been recently introduced from sonora into at least one home in la paz , baja california sur , mexico as pets , where it successfully reproduced ( patricia galina , personal communication to rwm ) . it likely occurs as introduced individuals or populations in north america and possibly elsewhere , although in this case many individuals are likely hybrids of gopherus morafkai x agassizii .
murray , roy c . ; schwalbe , cecil r . ; bailey , scott j . ; cuneo , s . peder ; hart , scott d . 1996 . reproduction in a population of the desert tortoise , gopherus agassizii , in the sonoran desert . herpetological natural history 4 ( 1 ) : 83 - 88
ennen , joshua r . ; jeffrey e . lovich , katherin p . meyer , curtis bjurlin , and terence r . arundel 2012 . nesting ecology of a population of gopherus agassizii at a utility - scale wind energy facility in southern california . copeia 2012 ( 2 ) : 222 - 228 . - get paper here
riedle , j . daren , roy c . averill - murray , clayton l . lutz and darren k . bolen . 2008 . habitat use by desert tortoises ( gopherus agassizii ) on alluvial fans in the sonoran desert , south - central arizona . copeia 2008 ( 2 ) : 414 - 420 - get paper here
lovich , jeffrey e . ; roy c . averill - murray , mickey agha , joshua r . ennen , and meaghan austin 2017 . variation in annual clutch phenology of sonoran desert tortoises ( gopherus morafkai ) in central arizona herpetologica dec 2017 , vol . 73 , no . 4 : 313 - 322 . - get paper here
tissue samples ( leg muscle ) were dissected from the lectotype of gopherus agassizii ( cooper , 1861 ) ( usnm 7888 ) and the holotype of gopherus lepidocephalus ( ottley & velázques solis , 1989 ) ( brigham young university [ byu ] 39706 ) . genomic dna was extracted from approximately 10 mg of tissue . the lectotype of gopherus agassizii was likely preserved in ethanol yet the holotype of gopherus lepidocephalus was initially well - preserved in formalin . subsequently , both specimens were stored in 70 % ethanol . to remove fixatives , tissues were washed twice in pbs , ph 7 . 2 ( 50 mm potassium phosphate , 150 mm nacl ) as recommended in the dna easy extraction kit ( qaigen ) for tissue exposed to formalin . subsequently , higher yields of dna were achieved using our standard extraction method , rather than the dna easy extraction kit , as follows : digestion of the tissue was carried out at 52 °c in a lysis buffer ( tris 6 . 06g , na2edta 0 . 93g , nacl 5 . 85g and sds 1 . 0g , 500ml ddh2o , ph 8 . 5 ) and spiked daily with 12 . 5 µl of proteinase k ( roche ) until the tissue sample was completely digested ( 5–7 days ) . purification used two washes with phenol : chloroform : isoamyl alcohol followed by a final wash of chloroform : isoamyl alcohol .
berry , kristin h . ; julie l . yee , ashley a . coble , william m . perry , and timothy a . shields 2013 . multiple factors affect a population of agassiz ' s desert tortoise ( gopherus agassizii ) in the northwestern mojave desert . herpetological monographs 27 ( 1 ) : 87 - 109 - get paper here
( a ) phylogenetic relationships and divergence times of three desert tortoise species . total rna was sequenced for three individuals per taxon and mapped to the gopherus agassizii assembly . ( b ) representation of gene ontology terms shared by genes with fixed unique single nucleotide polymorphisms in each species of desert tortoise . treemap boxes are sized according to uniqueness .
s9 dataset . biomart results for human ensembl gene id and gene ontology categories for accelerated genes with ka / ks > 1 between agassiz’s desert tortoise ( gopherus agassizii ) and western painted turtle ( chrysemys picta bellii ) , chinese softshell turtle ( pelodiscus sinensis ) , green sea turtle ( chelonia mydas ) , and chicken ( gallus gallus ) .
edwards 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 the gopherus morafkai–g . agassizii group . zookeys 562 : 131 - 158 . doi : 10 . 3897 / zookeys . 562 . 6124 - get paper here
aguirre g , morafka dj , adest ga ( 1997 ) conservation strategies for the bolson tortoise , gopherus flavomarginatus , in the chihuahuan desert . in : abbema jv , pritchard pch , editors . proceedings : conservation , restoration , and management of tortoises and turtles–an international conference . new york : new york turtle & tortoise society . pp . 333–338 .
the rounded ventral surface of the rear feet ( i–ii ) and multiple enlarged , raised scales present on surface of forelegs generally ( iii–iv ) diagnose gopherus evgoodei in relation to other species of desert tortoises . i–ii same individual in figure from rancho las cabras near alamos , sonora , mexico ( in tropical deciduous forest ) iii–iv two individuals from rancho las cabras .
scientific names consist of the genus ( generic name ) followed by a specific or species name . occasionally a third name is added to designate so - called sub - species , populations of the species with distinct characteristics . for more precision , the scientific name is often followed by the name of the person who first assigned the name and description . for example , the desert tortoise may be referred to as xerobates agassizii cooper . if the scientific name has undergone changes since the original description , then the describer ' s name is placed in parenthesis . for example , if the desert tortoise is in the genus gopherus then the name becomes gopherus agassizii ( cooper ) because cooper designated it as xerobates in his original description .
gopherus morafkai occurs in upland habitats in the sonoran desert scrub ( brown et al . 1979 ) with rocky outcrops and palo verde - saguaro cactus communities and ecotonal desert grasslands ( van devender 2002 ) . within these habitats , gopherus morafkai is generally found along rocky slopes , or bajadas , of desert mountain ranges , with breeding populations occurring as high as 1 , 420 m elevation and individual observation records occurring to 2 , 380 m ( flesch et al . 2010 ) . the species typically occupies excavated or eroded burrows underneath rocks or boulders . consequently , geology and resultant burrow availability among mountain ranges is an important determinant in regulating population density ( averill - murray et al . 2002a , b ) . low density populations of gopherus morafkai also occur along alluvial fans and in intermountain valleys , where individuals utilize desert washes and associated caliche caves as shelter sites ( riedle et al . 2008 ; grandmaison et al . 2010 ) . these peripheral populations provide important genetic linkages between disjunct mountain ranges ( edwards 2003 ; edwards et al . 2004 ; averill - murray and averill - murray 2005 ) .
sullivan , brian k . ; roy averill - murray , keith o . sullivan , justin r . sullivan , elizabeth a . sullivan , and j . daren riedle 2014 . winter activity of the sonoran desert tortoise ( gopherus morafkai ) in central arizona . chelonian conservation and biology jul 2014 , vol . 13 , no . 1 : 114 - 119 . - get paper here
mack , jeremy s . ; kristin h . berry , david m . miller , and andrea s . carlson 2015 . factors affecting the thermal environment of agassiz ' s desert tortoise ( gopherus agassizii ) cover sites in the central mojave desert during periods of temperature extremes journal of herpetology sep 2015 , vol . 49 , no . 3 : 405 - 414 . - get paper here
i ask because as soon as i announced having adopted a male gopherus , a guy i know through work connections posted on my facebook saying his family has a female
if i ever wanted to breed them
. i kinda wanted to slap the guy for suggesting it , what with so many existing gopherus in ca needing homes and the fact that captive hatchlings can ' t be returned to the wild . i want to say the adoption cttc adoption agent who helped us told me breeding them is illegal , but now i can ' t recall exactly . i already told the
match maker
no thanks , but i wondered if breeding them is frowned upon if i should tell him so he isn ' t throwing that offer around to anyone else and creating more homeless tortoises .
representation of gene ontology terms for biological processes shared by genes with pairwise ka / ks > 1 in comparisons of gopherus agassizii to ( a ) chrysemys picta bellii and ( b ) chelonia mydas . treemap boxes are both sized and colored by uniqueness . abbreviations in ( a ) : rb , receptor biosynthesis , lcfacb , long chain fatty acyl - coa biosynthesis , mtmd , membrane to membrane docking .
edwards , taylor ; mercy vaughn , philip c . rosen , cristina meléndez torres , alice e . karl , melanie culver and robert w . murphy 2016 . shaping species with ephemeral boundaries : the distribution and genetic structure of desert tortoise ( gopherus morafkai ) in the sonoran desert region . journal of biogeography43 ( 3 ) : 484–497 , doi : 10 . 1111 / jbi . 12664 - get paper here
murphy , robert w . ; kristin h . berry , taylor edwards , alan e . leviton , amy lathrop , j . daren riedle 2011 . 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 ; doi : 10 . 3897 / zookeys . 113 . 1353 - get paper here
using the alignment of murphy et al . ( 2007 ) , primers were designed for a 423 bp fragment that was diagnostic for haplotypes of gopherus agassizii . the forward primer gocytl ( 5’ - caattcgattcttcctagtagc - 3’ ) was located in the nadh3 gene and reverse primer gocyth ( 5’ - ggctgagaaggatagtattagtattgg - 3’ ) located on nd4 . attempts to amplify the holotype sample of gopherus lepidocephalus ( byu 39706 ) failed after numerous attempts using these two primers . because dna exposed to formalin is prone to degradation and fragmentation ( bucklin and allen 2004 ) , several internal primers were designed and used in various combinations until amplification was successful . eventually we amplified a 225 bp fragment using the original gocytl forward primer and a new internal reverse primer ( lepidond3h3 : 5’ - ttggtgtcattttgatagccgtgaag - 3’ ) that straddles the trnaarg and nd4l genes ; one bp was not confidently resolved .
a new species of desert tortoise has been discovered and described by a conservation geneticist with the university of arizona . the turtle , called the goode ' s thornscrub tortoise ( gopherus evgoodei ) was discovered in sinaloa , mexico , in thorn scrub and tropical deciduous forests . it is only found in these areas , according to ua geneticist taylor edwards and because of this , it has the smallest range of the three other known desert tortoises .
ottley and velázques solis ( 1989 ) described a new species , xerobates lepidocephalus , from the cape region of baja california sur , mexico . ecologically , the species occurs on sloped or hillside areas and it is not reported to live in burrows . this habitat choice closely resembles that of tortoises living in the sonoran desert , specifically those tortoises occurring east and south of the colorado river . these tortoises , called sonoran desert tortoises ( van devender 2002 ) , differ substantially from tortoises in the mojave desert . in general , sonoran tortoises live in rock crevices on steep slopes and hill tops ( riedle et al . 2008 ) and mojave desert tortoises live in burrows in valleys and on alluvial fans ( berry et al . 2002 ) . morphologically , gopherus lepidocephalus is most similar to tortoises on tiburon island off the coast of sonora , mexico and the species was considered to be a junior synonym of gopherus agassizii by crumly and grismer ( 1994 ) .


condica punctifera is a moth of the noctuidae family .

it is found in southern florida and on the bahamas , cuba , jamaica , hispaniola and puerto rico .

the wingspan is 27 mm .

adults are brown with a large reniform spot which is often accented with gray-white in the lower half .

there is also a black basal dash .

the hindwings are suffused with black-brown .

condica punctifera

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celaena punctifera walker , [ 1857 ] 1856 , list of the specimens of lepidopterous insects in the collection of the british museum , 10 : 263 .
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celaena semifurca walker , 1857 , list of the specimens of lepidopterous insects in the collection of the british museum , 11 : 732 .
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