Spalax: Difference between revisions

Spalax
	Greater blind mole-rat (S. microphthalmus)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Rodentia
Family:	Spalacidae
Subfamily:	Spalacinae
Genus:	Spalax; Güldenstädt, 1770
Type species
	Spalax microphthalmus; Güldenstädt, 1770
Species
	Spalax antiquus; Spalax arenarius; Spalax giganteus; Spalax graecus; Spalax istricus; Spalax microphthalmus; Spalax uralensis; Spalax zemni

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Spalax is a genus of rodent in the family Spalacidae, subfamily Spalacinae (blind mole-rats). It is one of two extant genera in the subfamily Spalacinae, alongside Nannospalax.^[2]

Species in this genus are found in eastern Europe and western & central Asia.^[3] They are completely blind and have a subterranean lifestyle.^[4]

Taxonomy

Prior to 2013, Spalax was widely considered the only member of Spalacinae, with all blind mole-rat species being grouped within it. However, phylogenetic and morphological evidence supported some of the species within it forming a distinct lineage that diverged from the others during the Late Miocene, when a marine barrier formed between Anatolia and the Balkans. These species were reclassified into the genus Nannospalax, making Spalax one of two extant spalacine genera.^[2]

Species of genus Spalax

Mehely's blind mole-rat, S. antiquus^[5]
Sandy blind mole-rat, S. arenarius
Giant blind mole-rat, S. giganteus
Bukovina blind mole-rat, S. graecus
Oltenia blind mole-rat, S. istricus (possibly extinct)^[5]
Greater blind mole-rat, S. microphthalmus
Kazakhstan blind mole-rat, S. uralensis
Podolsk blind mole-rat, S. zemni

References

^ Wilson, D. E.; Reeder, D. M., eds. (2005). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0. OCLC 62265494.
^ ^a ^b Hadid, Yarin; Németh, Attila; Snir, Sagi; Pavlíček, Tomáš; Csorba, Gábor; Kázmér, Miklós; Major, Ágnes; Mezhzherin, Sergey; Rusin, Mikhail; Coşkun, Yüksel; Nevo, Eviatar (2012-01-09). "Is Evolution of Blind Mole Rats Determined by Climate Oscillations?". PLOS ONE. 7 (1): e30043. Bibcode:2012PLoSO...730043H. doi:10.1371/journal.pone.0030043. ISSN 1932-6203. PMC 3253805. PMID 22253871.
^ Database, Mammal Diversity (2022-02-01), Mammal Diversity Database, doi:10.5281/zenodo.5945626, retrieved 2022-03-08
^ Lamarck, Jean-Baptiste (1809). Philosophie zoologique ou exposition des considérations relatives à l'histoire naturelle des animaux. Dentu.
^ ^a ^b Németh, Attila; Homonnay, Zalán G.; Krízsik, Virág; Csorba, Máté; Pavlíček, Tomáš; Hegyeli, Zsolt; Hadid, Yarin; Sugár, Szilárd; Farkas, János; Csorba, Gábor (2013). "Old views and new insights: taxonomic revision of the Bukovina blind mole rat, Spalax graecus (Rodentia: Spalacinae)" (PDF). Zoological Journal of the Linnean Society. 169 (4): 903–914. doi:10.1111/zoj.12081. S2CID 82892241.

Jansa, S. A.; Weksler, M. (2004). "Phylogeny of muroid rodents: relationships within and among major lineages as determined by IRBP gene sequences". Molecular Phylogenetics and Evolution. 31 (1): 256–76. doi:10.1016/j.ympev.2003.07.002. PMID 15019624.
Michaux, J.; Reyes, A.; Catzeflis, F. (2001). "Evolutionary history of the most speciose mammals: molecular phylogeny of muroid rodents". Molecular Biology and Evolution. 18 (11): 2017–2031. doi:10.1093/oxfordjournals.molbev.a003743. PMID 11606698.
Musser, G. G. and M. D. Carleton (2005). "Superfamily Muroidea." pp. 894–1531 in Wilson, D. E. and D. M. Reeder, eds. Mammal Species of the World: a Taxonomic and Geographic Reference. 3rd ed. Baltimore: Johns Hopkins University Press.
Norris, R. W.; Zhou, K. Y.; Zhou, C. Q.; Yang, G.; Kilpatrick, C. W.; Honeycutt, R. L. (2004). "The phylogenetic position of the zokors (Myospalacinae) and comments on the families of muroids (Rodentia)". Molecular Phylogenetics and Evolution. 31 (3): 972–978. doi:10.1016/j.ympev.2003.10.020. PMID 15120394.
Nowak, R. M. (1999). Walker's Mammals of the World, II. London: Johns Hopkins University Press ISBN 978-0-8018-5789-8
Steppan, S. J.; Adkins, R. A.; Anderson, J. (2004). "Phylogeny and divergence date estimates of rapid radiations in muroid rodents based on multiple nuclear genes". Systematic Biology. 53 (4): 533–553. doi:10.1080/10635150490468701. PMID 15371245.
Topachevskii, V. A. (1976) Fauna of the USSR. Volume III: Mammals. Issue 3: Mole rats, Spalacidae. New Delhi: Amerind.
Manov, Irena; Hirsh, Mark; Iancu, Theodore C; Malik, Assaf; Sotnichenko, Nick; Band, Mark; Avivi, Aaron; Shams, Imad (2013). "Pronounced cancer resistance in a subterranean rodent, the blind mole-rat, Spalax: in vivo and in vitro evidence". BMC Biology. 11: 91. doi:10.1186/1741-7007-11-91. PMC 3750378. PMID 23937926.

[msw3-1] Wilson, D. E.; Reeder, D. M., eds. (2005). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0. OCLC 62265494.

[:12-2] Hadid, Yarin; Németh, Attila; Snir, Sagi; Pavlíček, Tomáš; Csorba, Gábor; Kázmér, Miklós; Major, Ágnes; Mezhzherin, Sergey; Rusin, Mikhail; Coşkun, Yüksel; Nevo, Eviatar (2012-01-09). "Is Evolution of Blind Mole Rats Determined by Climate Oscillations?". PLOS ONE. 7 (1): e30043. Bibcode:2012PLoSO...730043H. doi:10.1371/journal.pone.0030043. ISSN 1932-6203. PMC 3253805. PMID 22253871.

[3] Database, Mammal Diversity (2022-02-01), Mammal Diversity Database, doi:10.5281/zenodo.5945626, retrieved 2022-03-08

[Lamarck2-4] Lamarck, Jean-Baptiste (1809). Philosophie zoologique ou exposition des considérations relatives à l'histoire naturelle des animaux. Dentu.

[ReferenceA-5] Németh, Attila; Homonnay, Zalán G.; Krízsik, Virág; Csorba, Máté; Pavlíček, Tomáš; Hegyeli, Zsolt; Hadid, Yarin; Sugár, Szilárd; Farkas, János; Csorba, Gábor (2013). "Old views and new insights: taxonomic revision of the Bukovina blind mole rat, Spalax graecus (Rodentia: Spalacinae)" (PDF). Zoological Journal of the Linnean Society. 169 (4): 903–914. doi:10.1111/zoj.12081. S2CID 82892241.

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[2]

[3]

[4]

[5]

@@ Line 2: / Line 2: @@
 {{for|the French record company|Spalax (record company)}}
 {{automatic taxobox
+| image = Філія ЛПЗ НАНУ "Стрільцівський степ" Spalax microphthalmus.jpg
-| fossil_range = {{fossil range|Early Miocene|Recent|Early [[Miocene]]–Recent}}
+| image_caption = [[Greater blind mole-rat]] (''S. microphthalmus'')
-| image = Palestine_Mole-rat_1.jpg
-| image_caption = [[Middle East blind mole-rat]], (''Spalax ehrenbergi'')
-| parent_authority = [[John Edward Gray|Gray]], 1821
 | taxon = Spalax
-| authority = [[Guldenstaedt]], 1770
+| authority = [[Johann Anton Güldenstädt|Güldenstädt]], 1770
+| type_species = ''[[Spalax microphthalmus]]''<ref name=msw3>{{MSW3|id=13000025}}</ref>
+| type_species_authority = [[Güldenstädt]], 1770
 | subdivision_ranks = [[Species]]
-| subdivision =
+| subdivision = ''[[Spalax antiquus]]''<br />
-''[[Spalax antiquus]]''<br />
 ''[[Spalax arenarius]]''<br />
-''[[Spalax carmeli]]''<br />
-''[[Spalax ehrenbergi]]''<br />
-''[[Spalax galili]]''<br />
 ''[[Spalax giganteus]]''<br />
-''[[Spalax golani]]''<br />
 ''[[Spalax graecus]]''<br />
 ''[[Spalax istricus]]''<br />
-''[[Spalax judaei]]''<br />
-''[[Spalax leucodon]]''<br />
 ''[[Spalax microphthalmus]]''<br />
-''[[Spalax munzuri]]''<br />
-''[[Spalax nehringi]]''<br />
 ''[[Spalax uralensis]]''<br />
 ''[[Spalax zemni]]''
-| synonyms = ''Nannospalax'' [[Theodore Sherman Palmer|Palmer]], 1903
 }}
+'''''Spalax''''' is a [[genus]] of [[rodent]] in the family [[Spalacidae]], subfamily [[Spalacinae]] (blind mole-rats). It is one of two extant genera in the subfamily [[Blind mole-rat|Spalacinae]], alongside ''[[Nannospalax]]''.''<ref name=":12">{{Cite journal |last1=Hadid |first1=Yarin |last2=Németh |first2=Attila |last3=Snir |first3=Sagi |last4=Pavlíček |first4=Tomáš |last5=Csorba |first5=Gábor |last6=Kázmér |first6=Miklós |last7=Major |first7=Ágnes |last8=Mezhzherin |first8=Sergey |last9=Rusin |first9=Mikhail |last10=Coşkun |first10=Yüksel |last11=Nevo |first11=Eviatar |date=2012-01-09 |title=Is Evolution of Blind Mole Rats Determined by Climate Oscillations? |journal=PLOS ONE |language=en |volume=7 |issue=1 |pages=e30043 |doi=10.1371/journal.pone.0030043 |issn=1932-6203 |pmc=3253805 |pmid=22253871|bibcode=2012PLoSO...730043H |doi-access=free }}</ref>''
-The genus '''''Spalax''''' contains the '''blind''', '''fossorial''', or '''subterranean mole rats''', which are one of several types of [[rodent]]s that are called "mole rats".<ref name=EoM>{{cite book|editor= Macdonald, D.|year= 2006|title= The Encyclopedia of Mammals|publisher= Oxford University Books|location= Oxford|page= [https://archive.org/details/encyclopediaofma00mals_0/page/203 203]|isbn= 978-0-87196-871-5|url-access= registration|url= https://archive.org/details/encyclopediaofma00mals_0/page/203}}</ref> The [[Hystricognathi|hystricognath]] mole-rats of the family [[Bathyergidae]] are completely unrelated, but some other forms are also in the family [[Spalacidae]]. [[Zokor]]s (subfamily [[Myospalacinae]]), root rats, and [[bamboo rat]]s (subfamily [[Rhizomyinae]]) are spalacids sometimes referred to as mole rats.  Blind mole rats are in the [[family (biology)|family]] [[Spalacidae]], but are unique enough to be given a separate [[subfamily]], '''Spalacinae'''. Alternate opinions on [[Taxonomy (biology)|taxonomy]] consider the blind mole-rats to be the only members of the family [[Spalacidae]] and rank other spalacid subfamilies as full families.  Other authors group all members of the [[Taxonomic rank|superfamily]] [[Muroidea]] into a single family, [[Muridae]]. The Spalacinae contains two [[genus|genera]] and eight [[species]].  Some authorities treat all species as belonging to a single genus, ''[[Spalax (genus)|Spalax]]''.
+Species in this genus are found in [[eastern Europe]] and [[Western Asia|western]] & [[central Asia]].<ref>{{Citation |last=Database |first=Mammal Diversity |title=Mammal Diversity Database |date=2022-02-01 |doi=10.5281/zenodo.5945626 |url=https://zenodo.org/record/5945626 |access-date=2022-03-08}}</ref> They are completely blind and have a subterranean lifestyle.<ref name="Lamarck2">{{cite book |author=Lamarck, Jean-Baptiste |url=https://archive.org/details/bub_gb_8NsTAAAAQAAJ |title=Philosophie zoologique ou exposition des considérations relatives à l'histoire naturelle des animaux |publisher=Dentu |year=1809}}</ref>
-''Spalax'' mole rats are truly [[Blind animals|blind]].<ref name=Lamarck>{{cite book|author=Lamarck, Jean-Baptiste |year=1809 |title=Philosophie zoologique ou exposition des considérations relatives à l'histoire naturelle des animaux|publisher=Dentu |url=https://archive.org/details/bub_gb_8NsTAAAAQAAJ }}</ref>  Their very small eyes are completely covered by a layer of skin. Unlike many other [[fossorial]] [[rodent]]s, ''Spalax'' mole rats do not have enlarged front claws and do not appear to use their forearms as a primary digging tool. Digging is almost exclusively conducted using their powerful [[incisor|front teeth]], which are separated from the rest of the mouth by a flap of skin. When a ''Spalax'' mole rat closes its mouth, its incisors are still on the outside. Blind mole rats may have evolved from [[Spalacidae|spalacids]] that used their front limbs to dig, because their [[olecranon process]]es are large relative to the rest of their arms.  The olecranon process is a part of the [[ulna]] bone where muscles attach, and [[fossorial|digging animals]] tend to have enlarged olecranon processes to provide a large surface for their large and powerful [[muscle]]s to attach.
+== Taxonomy ==
-Because they are completely blind, blind mole rats have been important laboratory animals in tests on how [[eye]]s and eye [[protein]]s function. Although Spalax has only atrophied subcutaneous eyes and it is sightless, its circadian rhythm is kept. A few publications (Avivi et al.) have proved that the circadian genes that control the biological clock are expressed in a similar manner as in sighted, above-ground mammals.
+Prior to 2013, ''Spalax'' was widely considered the only member of Spalacinae, with all blind mole-rat species being grouped within it. However, phylogenetic and morphological evidence supported some of the species within it forming a distinct lineage that diverged from the others during the [[Late Miocene]], when a marine barrier formed between [[Anatolia]] and the [[Balkans]]. These species were reclassified into the genus ''[[Nannospalax]]'', making ''Spalax'' one of two extant spalacine genera.''<ref name=":12"/>''
+==Species of genus ''Spalax'' ==
-== Resistance to cancer ==
+*[[Mehely's blind mole-rat]]'', S. antiquus''<ref name="ReferenceA">{{cite journal|title=Old views and new insights: taxonomic revision of the Bukovina blind mole rat, Spalax graecus (Rodentia: Spalacinae)|date=2013|doi=10.1111/zoj.12081|url=http://real.mtak.hu/7751/7/NemethA%202013%20zoj_12081%20Copy.pdf|last1=Németh |first1=Attila |last2=Homonnay |first2=Zalán G. |last3=Krízsik |first3=Virág |last4=Csorba |first4=Máté |last5=Pavlíček |first5=Tomáš |last6=Hegyeli |first6=Zsolt |last7=Hadid |first7=Yarin |last8=Sugár |first8=Szilárd |last9=Farkas |first9=János |last10=Csorba |first10=Gábor |journal=Zoological Journal of the Linnean Society |volume=169 |issue=4 |pages=903–914 |s2cid=82892241 }}</ref>
-Studies on the growth of [[fibroblast]]s ''[[in vitro]]'' of ''[[Spalax judaei]]'' and ''[[Spalax golani]]'' showed that the process of [[necrosis]] replaces the role of the systematic [[apoptosis]] normally used in most organisms. Generally low-oxygen conditions, such as those common in blind mole rats’ burrows, usually cause cells to undergo apoptosis. One study showed that in adaptation to a higher tendency of cell death, blind mole rats evolved a mutation in the [[tumor suppressor]] protein [[p53]], also used in humans, to prevent cells from undergoing apoptosis.<ref>{{cite journal|last1=Avivi|first1=A|last2=Ashur-Fabian|first2=O|last3=Joel|first3=A|last4=Trakhtenbrot|first4=L|last5=Adamsky|first5=K|last6=Goldstein|first6=I|last7=Amariglio|first7=N|last8=Rechavi|first8=G|last9=Nevo|first9=E|title=P53 in blind subterranean mole rats – loss-of-function versus gain-of-function activities on newly cloned Spalax target genes|journal=Oncogene|date=16 October 2006|volume=26|issue=17|pages=2507–2512|doi=10.1038/sj.onc.1210045|pmid=17043642|doi-access=free}}</ref> Human cancer patients have similar mutations, and blind mole rats were thought to be more susceptible to cancer because their cells cannot undergo apoptosis. However, after a specific amount of time (within 3 days according to one study), the cells in blind mole rats release [[interferon-beta]] (which the immune system normally uses to counter viruses) in response to over-proliferation of cells caused by the suppression of apoptosis. In this case, the interferon-beta triggers cells to undergo necrosis, and this mechanism also kills cancer cells in blind mole rats. Because of tumor suppression mechanisms such as this, blind mole rats and other spalacids are resistant to cancer.<ref name="Gorbunova">{{cite journal|last=Saey|first=Tina Hesman|title=Cancer cells self-destruct in blind mole rats|url=http://www.sciencenews.org/view/generic/id/346267/description/Cancer_cells_self-destruct_in_blind_mole_rats|journal=[[Science News]]|accessdate=27 November 2012|date=5 November 2012}}</ref><ref>{{Cite journal | last1 = Gorbunova | first1 = V. | last2 = Hine | first2 = C. | last3 = Tian | first3 = X. | last4 = Ablaeva | first4 = J. | last5 = Gudkov | first5 = A. V. | last6 = Nevo | first6 = E. | last7 = Seluanov | first7 = A. | doi = 10.1073/pnas.1217211109 | title = Cancer resistance in the blind mole rat is mediated by concerted necrotic cell death mechanism | journal = Proceedings of the National Academy of Sciences | volume = 109 | issue = 47 | pages = 19392–6 | year = 2012 | pmid =  23129611| pmc = 3511137| bibcode = 2012PNAS..10919392G | doi-access = free }} Retrieved 27 November 2012</ref><ref>{{cite conference|url=https://pag.confex.com/pag/xxii/webprogram/Paper10495.html|title=Fighting Cancer Underground: Hypoxia Tolerant Spalax Hides the Key for Treatment|date=2014|first1=Imad|last1=Shams|first2=Irena|last2=Manov|first3=Assaf|last3=Malik|first4=Mark|last4=Band|first5=Aaron|last5=Avivi|conference=The Plant & Animal Genome XXII Conference}}</ref>
+*[[Sandy blind mole-rat]], ''S. arenarius''
+*[[Giant blind mole-rat]], ''S. giganteus''
-The involvement of interferon in the so-called concerted cell death of Spalax cells via necrosis was highly criticized.<ref name="Gorbunova" /> Serious questions have been raised on the inconsistent methodology used that led to this speculation.
+*[[Bukovina blind mole-rat]], ''S. graecus''
+*[[Oltenia blind mole-rat]], ''S. istricus'' (possibly [[Extinction|extinct]])<ref name="ReferenceA" />
-==Classification==
+*[[Greater blind mole-rat]], ''S. microphthalmus''
+*[[Kazakhstan blind mole-rat]], ''S. uralensis''
-'''Subfamily Spalacinae'''
+*[[Podolsk blind mole-rat]], ''S. zemni''
-*Genus ''Spalax'' – Blind mole rats
-**''[[Spalax antiquus]]''<ref name="ReferenceA">{{cite conference|title=Old views and new insights: taxonomic revision of the Bukovina blind mole rat, Spalax graecus (Rodentia: Spalacinae)|date=2013|doi=10.1111/zoj.12081|url=http://real.mtak.hu/7751/7/NemethA%202013%20zoj_12081%20Copy.pdf}}</ref>
-**[[Sandy mole rat]], ''Spalax arenarius''
-**[[Mount Carmel blind mole rat]], ''Spalax carmeli''
-**[[Middle East blind mole rat]], ''Spalax ehrenbergi''
-**[[Upper Galilee Mountains blind mole rat]], ''Spalax galili''
-**[[Giant mole rat]], ''Spalax giganteus''
-**[[Golan Heights blind mole rat]], ''Spalax golani''
-**[[Balkan mole rat]], ''Spalax graecus''
-**''[[Spalax istricus]]''<ref name="ReferenceA"/>
-**[[Judean Mountains blind mole rat]], ''Spalax judaei''
-**[[Lesser mole rat]], ''Spalax leucodon''
-**[[Greater mole rat]], ''Spalax microphthalmus''
-**[[Munzur mole rat]], ''Spalax munzuri''
-**[[Nehring's blind mole rat]], ''Spalax nehringi''
-**[[Kazakhstan blind mole rat]], ''Spalax uralensis''
-**[[Podolsk mole rat]], ''Spalax zemni''
 ==References==
 {{Reflist|2}}
-^ Irena Manov, Mark Hirsh, Theodore C Iancu, Assaf Malik, Nick Sotnichenko, Mark Band, Aaron Avivi and Imad Shams† (2013) Pronounced cancer resistance in a subterranean rodent, the blind mole-rat, Spalax: in vivo and in vitro evidence. BMC Biology [https://bmcbiol.biomedcentral.com/articles/10.1186/1741-7007-11-91]
 ==Further reading==
@@ Line 74: / Line 48: @@
 * {{cite journal | last1 = Steppan | first1 = S. J. | last2 = Adkins | first2 = R. A. | last3 = Anderson | first3 = J. | year = 2004 | title = Phylogeny and divergence date estimates of rapid radiations in muroid rodents based on multiple nuclear genes | journal = Systematic Biology | volume = 53 | issue = 4 | pages = 533–553 | doi = 10.1080/10635150490468701 | pmid = 15371245 }}
 * Topachevskii, V. A. (1976) ''Fauna of the USSR. Volume III: Mammals. Issue 3: Mole rats, Spalacidae''. New Delhi: Amerind.
+*{{cite journal | first1=Irena |last1=Manov |first2=Mark |last2=Hirsh |first3=Theodore C |last3=Iancu |first4=Assaf |last4=Malik |first5=Nick |last5=Sotnichenko |first6=Mark |last6=Band |first7=Aaron |last7=Avivi |first8=Imad |last8=Shams |year=2013 |title=Pronounced cancer resistance in a subterranean rodent, the blind mole-rat, Spalax: in vivo and in vitro evidence|journal=BMC Biology|volume=11 |page=91 |doi=10.1186/1741-7007-11-91 |pmid=23937926 |pmc=3750378 |doi-access=free }}
 {{Spalacidae nav}}
+{{Myomorpha|M.|state=collapsed}}
 {{Taxonbar|from=Q951670}}
-[[Category:Spalax| ]]
+[[Category:Spalax ]]
 [[Category:Rodent genera]]
 [[Category:Extant Miocene first appearances]]
-[[Category:Taxa named by John Edward Gray]]
+[[Category:Taxa named by Johann Anton Güldenstädt]]
-[[ar:خلد]]
 [[fr:Spalacinae]]
 [[pl:Ślepce]]