Cytogenetic analysis of cryptic species in allopatric population (2n = 60, NF = 82) of Nannospalax sp. (Spalacidae, Rodentia): conventional karyotype, constitutive heterochromatin, and NOR localization

Cytogenetic analysis of cryptic species in allopatric population (2n = 60, NF = 82) of Nannospalax sp. (Spalacidae, Rodentia): conventional karyotype, constitutive heterochromatin, and NOR localization | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Cytogenetic analysis of cryptic species in allopatric population (2n = 60, NF = 82) of Nannospalax sp. (Spalacidae, Rodentia): conventional karyotype, constitutive heterochromatin, and NOR localization Gökhan YÜRÜMEZ, Yüksel COŞKUN This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6277258/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 08 Jan, 2026 Read the published version in Biologia → Version 1 posted 6 You are reading this latest preprint version Abstract In the present study, the 2n = 60, NF = 82 chromosomal form of blind mole rats, Nannospalax sp. , from Arapgir-Malatya Province in Turkey was investigated. Conventional chromosome staining, NOR (Nucleolus Organizer Region), and C-banding analysis were carried out on specimens of this population. The karyotype including 10 submetacentric pairs and 19 acrocentric pairs of autosomes and fundamental number is (NFa) 78. In all of the specimens studied the NOR’s were localized in the telomeric regions of the short arms of the 2 autosome pairs (nos. 2 and 7), and only one homologue of the pairs nos. 4 and 10 autosomes has a positive silver signal. C-heterochromatin regions were found in the centromeric region of whole bi-armed autosomal pairs. One of the X chromosomes has a pericentromeric C-positive band, whereas the other has a dot-like C-positive band. Nannospalax mole rat karyotype Malatya Turkey Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Mole rats (family Spalacidae) are the major component of steppe habitats in the Palearctic region (Topachevski 1976; Savic and Soldatovic 1977 ), and is the most speciation groups taxon, which is currently regarded to be composed of more than 60 chromosomal forms in 2 genera ( Spalax and Nannospalax ) in distributon area and the genus Nannospalax includes three morphospecies: Nannospalax leucodon , Nannospalax nehringi , and Nannospalax ehrenbergi in Turkey (Ellerman and Morrison-Scoot 1951; Topachevski 1976; Musser and Carleton 2005 ). Despite almost 30–40 million years of the family Spalacidae evolutionary background, speciation is still in progress and many populations are at different levels of the process (Savic and Nevo 1990 ; Nevo et al. 1995 ; Ünay 1996 ; Şen and Sarica 2011). The genus Nannospalax which is one of the youngest genera (about 24 million years ago) of this family (Hadid et al. 2012 ), shows a relatively high rate of karyotypic rearrangements and molecular divergence among mammals. These evolutionary dynamics provoke taxonomic uncertainties on the one hand but make this group a perfect model for evolutionary research on the other hand (Savic and Nevo 1990 ; Nevo et al. 1995 ; He et al. 2019 ). Overall, the Nannospalax mole rats abound with cryptic species (Nevo et al. 2001 ; Csorba et al. 2015 ; Savic et al. 2017; Bugarski-Stanojević at al. 2022, 2024). Cryptic species are those that cannot readily be distinguished on the basis of phenotypic variation alone (Struck et al. 2018 ); they have created challenges in myriad areas of the life sciences, especially in taxonomic systems. Not all “cryptic” species, however, are truly cryptic; many are simply underexplored morphologically (Nastasi et al. 2024 ). Cryptic species complexes where there is a large diversity of karyomorphs with autapomorphies must have originated due to plasticity in the ancestral karyomorph. The multiple karyotypic forms found in cryptic species could constitute, therefore, important examples of non-adaptive or adaptive chromosomal radiation, depending on the relation of the character with the environment (Kavalco and Pasa 2023 ). Chromosomal studies have often significantly contributed to understanding of the phylogeny and systematics of small mammals owing to extensive karyotypic variation recorded in their populations (Zima 2000 ; Arslan et al. 2011a ). Changes in C-heterochromatin amount and distribution are among important mechanisms producing this variation. C- and Ag-NOR banding techniques are frequently used in animals and thus are useful for examining intra- and interspecific chromosomal differences between closely related species (Arslan et al. 2011a ; Zhu et al. 2019 ). The structure, number, and morphology of the nucleolar organizer region may be specific to populations, species, and subspecies. Nucleolus Organizer Regions (NORs) are frequently used as markers to show intra- and inter-species chromosomal polymorphism in animals. Species with limited gene exchange owing to geographic isolation have high NORs and chromosomal diversity. Therefore, even in small but isolated populations of these species, different karyotypes are found. The homogeneity of this character and the degree of diversity within a taxon largely determines the use of NORs in explaining relationships (Yüksel and Gaffaroğlu 2008 ). Conventional karyological studies carried out in Malatya province: Yüksel ( 1984 ) 2n = 60, NF = 80 from Yazıhan; Gülkaç and Yüksel ( 1989 ) 2n = 60, NF = 80 from Akçadağ and NF = 82 from Arguvan; Nevo et al. ( 1995 ) 2n = 60, NF = 78 from 30 km west of Malatya; Ivanitskaya et al. ( 1997 ) 2n = 60, NF = 78 from 12 km east of Malatya; Ulutürk et al. ( 2009 ) 2n = 60, NF = 82 from Arguvan, Hekimhan, and Arapgir; Coşkun et al. ( 2010 ) 2n = 60, NF = 78 from Doğanşehir, Kale and Akçadağ (Table 1 ). So far C-, and NOR banding of different chromosomal forms of Nannospalax 2n = 60 were achieved from Turkey by various authors (Ivanitskaya et al. 1997 , 2008 ; Gülkaç and Küçükdumlu 1999 ; Arslan and Bölükbaş 2010 ; Arslan et al. 2011b ; Aşan Baydemir et al. 2013 ; Yağcı 2017 ). However, there is not any banded karyotypes of Nannospalax population (2n = 60, NF = 82) in Arapgir (Malatya) region. The aim of the present study is to describe in detail the conventional chromosome staining and the distribution of the NORs and C-heterochromatin in the karyotype of Nannospalax sp. from Arapgir (Malatya). Material and methods Cytogenetic analyses were performed on three specimens of mole rat from Arapgir, Malatya (Fig. 1 ). Karyotype preparations were obtained from the bone marrow of animals treated with colchicine (Ford and Hamerton 1956 ). After preparation of chromosome slides, air-dried preparations were stained conventionally with Giemsa. Constitutive heterochromatin and nucleolus organizer regions were detected in individual autosomal and sex chromosome pairs via C-banding (Sumner, 1972 ), and Ag-NOR staining (Howell and Black 1980 ). From each specimen slides were prepared, and well-spread metaphase plates were analysed. The results were compared with previously published accounts on 2n = 60 chromosomal forms carried out in Turkey. Standard voucher specimens (skins and skulls) are deposited in the Department of Biology, Faculty of Science, Dicle University, Diyarbakır, and Batman University Science and Art Faculty, Biology Department, Turkey. Results Conventional karyotype All karyotyped mole rats from the population of Arapgir had identical karyotypes with 2n = 60, NF = 82 and NFa = 78. The autosomal set consisted of ten pairs submetacentric (chromosome nos. 1–10) and 19 pairs large-to-small acrocentric pairs (chromosome nos.11–29) of gradually decreasing size). The X chromosome was the second large submetacentric, and the Y chromosome was dot-sized acrocentric (Fig. 2 ). NOR banding patterns NORs were found in two bi-armed autosomal pairs (pairs 2 and 7) in the complements of all the specimens, and in the telomeric region of the short arms of those pairs, and only one homologue of the pairs numbers 4 and 10 autosomes has determined a positive silver signal (Fig. 3 ). C-banding In mole rats of Arapgir, C-positive heterochromatic blocks were located in centromeric region of all biarmed autosomes (1–10) and while the one of the X chromosomes has a large pericentromeric C-positive region, the other one X chromosome has a dot like centromeric C positive (Fig. 4 ) The distribution of C-heterochromatin followed the same pattern in all examined specimen sets. Some biarmed autosome pairs (numbers 1, 5, 7, and 8) exhibited distinct dark centromeric C-bands. Other biarmed autosome pairs (numbers 2, 3, 4, 6, 9, and 10) exhibited a weaker C-positive band. Discussion The 2n = 60 chromosomal form shows the largest distribution areas among chromosomal forms of Nannospalax in Anatolia. The 2n = 60 chromosomal form in Turkey were first recorded by Yüksel ( 1984 ) from Malatya Province in the central-eastern Anatolia. Later, 2n = 60, the most common chromosomal form of mole rats in central Anatolia, had been reported by numerous authors (Gülkaç and Yüksel 1989 ; Nevo et al. 1995 ; Ivanitskaya et al. 1997 ; Gülkaç and Küçükdumlu 1999 ; Ulutürk et al. 2009 ; Coşkun et al. 2010 , and see Table 1 ). The karyotype in the 2n = 60 chromosomal forms are variable and NF range of 72–84 was reported by previous authors (Matur and Sözen 2005 ; Kankılıç et al. 2007 , 2009 , 2010 ; Sözen et al. 2011 , 2013 ; Arslan et al. 2016 ). However, the geographical continuity of the known populations is in doubt. In the Malatya province two different chromosomal forms of 2n = 60 (2n = 60a, NF = 82 and 2n = 60b, NF = 78) reported by Coşkun et al. ( 2010 ), and they claimed that these two forms seem to be separated by the Tohma Stream. The NORs of Nannospalax populations in Malatya province were first studied by Ivanitskaya et al. ( 1997 ). According to these researchers, in all the samples, the NORs were localised in the short arms of 3 pairs of sub-telocentric chromosomes in this population. Gülkaç and Küçükdumlu ( 1999 ) determined that Nannospalax ( Spalax leucodon ) specimens obtained from Malatya, the karyotype consitsts of 30 chromosome pairs and the NORs were terminally located on 3 pairs of large subtelocentrics. All of the individuals displayed a consistent number of NORs per cell. NORs were not found on the sex chromosomes of Nannospalax ( Spalax leucodon ). Table 1 Karyotype records of the 2n = 60 chromosomal forms of Nannospalax from Turkey. NF – fundamental number of chromosomal arms (See Fig. 1 for location of the sites). 2n NF m/sm/st a X Y C band NOR Locality Reference 2n = 60 80 18 40 sm st -- -- Malatya Yazıhan Yüksel 1984 80 18 40 sm st -- -- Malatya Akçadağ Gülkaç and Yüksel 1989 82 20 38 sm a -- -- Arguvan 78 16 42 sm a -- -- Malatya 30 km W Nevo at al. 1995 78 16 42 -- -- 14 m/sm + 8a + XY 6 m/sm Malatya 12 km E Ivanitskaya et al. 1997 -- -- -- -- -- -- 6 m/sm Malatya Gülkaç and Küçükdumlu 1999 78 16 42 Sm a 16 + XY 8 m/sm (W) Çankırı, Orta, Sanabozu Ivanitskaya et al. 2008 9 m/sm (R) Bursa İnegöl 82 20 38 M a -- -- Malatya Arguvan, Hekimhan, Arapgir Ulutürk et al. 2009 Sm a -- -- Elazığ, Keban (Denizli) Coşkun et al. 2010 Erzincan, Çitköy, Dutluca Sivas Divriği 78 16 42 sm a -- -- Elazığ, Keban (Çirkan) Malatya Doğanşehir, Kale, Akçadağ 76 14 44 sm 14 m/sm + XY 8 m/sm Aksaray Gülağaç, Ortaköy, Ağaçören Arslan and Bölükbaş 2010 78 16 42 sm m 16 m/sm + XY Güzelyurt, Sarıyahşi, Merkez 74 12 46 Sm st 12 m/sm + XY 8 m/sm Konya Hadim, Karatay Arslan et al. 2011b 78 16 42 -- -- 16m/sm + XY + 10a 8 m/sm Ilgın, Höyük, Sarayönü 79 17 21– Sm st 17 + XY + 12a 8 m/sm Bozkır, Çumra, Selçuklu, Güneysınır, Meram 80 18 40 Sm St 18 + XY 8 m/sm Cihanbeyli 80 18 40 Sm a -- 8 m/sm Konya Cihanbeyli, Kulu Aşan Baydemir et al. 2013 -- -- -- -- -- 18 m/sm + XY 6 m/sm Bilecik Söğüt, Pazaryeri Yağcı 2017 82 20 38 Sm a 20 m/sm + XX 6 m/sm Malatya Arapgir This study Nevo et al. ( 1995 )d lkaç and Küçükdumlu (1999) reported that the NORs on homologue chromosomes do not exhibit heteromorphism in Nannospalax ( Spalax leucodon ). The number of NORs carrying chromosomes and the location of the NORs on the chromosomes usually show remarkable variation among species. Sözen et al. ( 2006 ) reported that it appears that changes in NORs have occurred in strict association with differentiation and evolution of species. Ivanitskaya et al. ( 2008 ) reported that both 2n = 60R and 2n = 60W chromosomal forms have NORs localized in the five pairs of sub-telocentric autosomes. Arslan et al. ( 2011b ) reported the NORs have in 4 pair autosomes of the four chromosomal forms (NF = 74, 78, 79 and 80) of 2n = 60 from Konya Distributions of NORs patterns were very similar to those in Malatya (Ivanitskaya et al. 1997 ; Gülkaç and Küçükdumlu 1999 ) and Bilecik (Yağcı 2017 ), but some differences in Bursa, Çankırı (Ivanitskaya et al. 2008 ), Aksaray (Arslan and Bölükbaş 2010 ), Konya (Arslan et al. 2011b ). Nannospalax ( S. leucodon ) samples (2n = 60, NF = 78) from Malatya, all subtelocentric chromosomes possess completely heterochromatic short arms, with the exception of the pair 7; among the acrocentric chromosomes only four pair; have pericentromeric blocks of heterochromatin; in the submetacentric X-chromosomes the pericentromeric blocks of heterochromatin are not as dark as in the autosomes (Ivanitskaya et al. 1997 ). Chromosomes of blind mole rats from Malatya (2n = 60, NF = 78, NFa = 74) given by Ivanitskaya et al. 1997 differed distinctly with the C-banding pattern of Arapgir population (2n = 60, NF = 82, NFa = 78). In terms of biarmed chromosomes, C-heterochromatin distributions were very similar those in Çankırı and Bursa (Ivanitskaya et al. 2008 ), Aksaray (Arslan and Bölükbaş 2010 ) and Konya (Arslan et al. 2011b ), but some differences in Malatya (Ivanitskaya et al. 1997 ). According to the acrocentric chromosomes C-heterochromatin distributions differ from in Malatya (Ivanitskaya et al. 1997 ) and Konya (Ilgın, Höyük, Sarayönü, Bozkır, Çumra, Selçuklu, Güneysınır and Meram) (Arslan et al. 2011b ). All sex chromosomes possesses C-heterochromatin blocks in all 2n = 60 populations in Anatolia (See Table 1 ). Nucleolus organizer regions and C-heterochromatin distributions have been detected for the first time in this study for Nannospalax sp. 2n = 60, NF = 82 chromosomal form of Turkey. To date, no detailed cytogenetic investigations have been performed on representatives of Nannospalax sp. which inhabit a small territory in the northern part of the Tohma River basin in Arapgir (Malatya). Coşkun et al ( 2010 ) stated that the two cryptic species (2n = 60a, NF = 82 and 2n = 60b, NF = 78) of mole rats have distributed in Malatya province and they argued the Tohma River acts as a barrier between two cryptic chromosomal forms. In this study, the presence of chromosomal forms 2n = 60a, NF = 82 ( Nannospalax sp. ) was confirmed. Morphological examination of specimens from both chromosomal forms is necessary to reveal morphological differences and, at last to a formal species description on this cryptic species. Our study demonstrates the uncovering hidden biodiversity. We, therefore, classified these mole rats as Nannospalax sp. but increasing the sample size is needed to rule out of this cryptic species. Declarations Funding The authors declare that no funds, grants, or other support was received during the preparation of this manuscript. Conflict of interest The authors declare no conflict of interest. Ethical approval No ethical approval is required, not applicable. Author contribution All authors made equal contributions to this study. Data availability All data supporting the findings of this study are available within the paper. References Arslan A, Bölükbaş F (2010) C-heterochromatin and NORs distrubution of mole rat, Nannospalax xanthodon from Aksaray. Turk Caryologia 63:398404. https://doi.org/10.1080/00087114.2010.10589752 Arslan A, Toyran K, Gözütok S, Yorulmaz T (2011a) C- and NOR stained karyotypes of mole rat, Nannospalax xanthodon (2n = 54) from Kırıkkale. 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Mol Cytogenet 12:41. https://doi.org/10.1186/s13039-019-0453-1 Zima J (2000) Chromosomal evolution in small mammals (Insectivora, Chiroptera, Rodentia). Hystrix It J Mamm 11:5–15. https://doi.org/10.4404/hystrix-11.2-4143 Cite Share Download PDF Status: Published Journal Publication published 08 Jan, 2026 Read the published version in Biologia → Version 1 posted Editorial decision: Major revisions 26 Jul, 2025 Reviewers agreed at journal 24 Jun, 2025 Reviewers invited by journal 24 Jun, 2025 Editor invited by journal 08 Jun, 2025 Editor assigned by journal 25 Mar, 2025 First submitted to journal 21 Mar, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6277258","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":475659204,"identity":"05d2f417-f088-4e63-a65c-943eeb789ee8","order_by":0,"name":"Gökhan YÜRÜMEZ","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA8UlEQVRIiWNgGAWjYFACNjZkng0QMzYeIEVLGkhLA0laDoNJvFrk3dvSHvzcwZC44fYZ0w0f/py3W9t+GGhLjU00Li2GZ44dN+w9A9RyLsfs5gye28nbziQCtRxLy23ApWVGepsEbxtQyxkes9s8EreTzQ4AtTA2HMarRfIvTMsfg3PJZucf4tciL5F2TBpuC0PCATuzGwRsMeA5liYt2yZhPPMMW9nNngPJCWY3gLYk4PGLfHubmeTbNhvZvjPM2278+GNnb3Y+/eGDDzU2uG05AKYkHGEKEsGMBBzKwbZAldrDBOxxKBwFo2AUjIIRDADiTmZYb9kikQAAAABJRU5ErkJggg==","orcid":"https://orcid.org/0000-0001-5415-2278","institution":"Batman University: Batman Universitesi","correspondingAuthor":true,"prefix":"","firstName":"Gökhan","middleName":"","lastName":"YÜRÜMEZ","suffix":""},{"id":475659205,"identity":"7b04768d-23b5-4447-95dd-a013d0423eb7","order_by":1,"name":"Yüksel COŞKUN","email":"","orcid":"","institution":"Dicle University: Dicle Universitesi","correspondingAuthor":false,"prefix":"","firstName":"Yüksel","middleName":"","lastName":"COŞKUN","suffix":""}],"badges":[],"createdAt":"2025-03-21 11:39:40","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6277258/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6277258/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s11756-025-02124-x","type":"published","date":"2026-01-08T15:57:22+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":85497024,"identity":"c8092182-6982-43e3-a1d2-128d01685492","added_by":"auto","created_at":"2025-06-26 14:03:43","extension":"jpeg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":255787,"visible":true,"origin":"","legend":"\u003cp\u003eStudy area and previous studies on \u003cem\u003eNannospalax sp.\u003c/em\u003epopulation from Malatya province (Also see table 1).\u003c/p\u003e","description":"","filename":"floatimage1.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-6277258/v1/80fa995dd1031c06b43d855d.jpeg"},{"id":85498116,"identity":"2969b351-20af-4e1a-a3d1-6ad26cc5245e","added_by":"auto","created_at":"2025-06-26 14:11:43","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":181399,"visible":true,"origin":"","legend":"\u003cp\u003eConventional karyotypes of \u003cem\u003eNannospalax sp. \u003c/em\u003e(Arapgir-Malatya).\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-6277258/v1/84c79c0ddf4d42a89e4855ee.png"},{"id":85497029,"identity":"6f0b225d-60e4-486e-ac1f-a2243e7c3b37","added_by":"auto","created_at":"2025-06-26 14:03:43","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":194758,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eS\u003c/strong\u003eilver nitrate stained (Ag-NOR) karyotypes of \u003cem\u003eNannospalax sp. \u003c/em\u003e(Arapgir-Malatya) (Black arrows indicate localization of NORs).\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-6277258/v1/2563ffada4088bbf59889cd1.png"},{"id":85498922,"identity":"d206d43a-9fa5-4323-8409-e68eee4dc888","added_by":"auto","created_at":"2025-06-26 14:19:43","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":233859,"visible":true,"origin":"","legend":"\u003cp\u003eC-banded karyotypes of \u003cem\u003eNannospalax sp. \u003c/em\u003e(Arapgir-Malatya) (Black arrows indicate localization of C-heterochromatin).\u003c/p\u003e","description":"","filename":"floatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-6277258/v1/e8a075a474360f13e85de117.png"},{"id":100069102,"identity":"c0735b33-5597-4db2-b08f-797de151a646","added_by":"auto","created_at":"2026-01-12 16:09:30","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1662117,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6277258/v1/c7559915-6154-4685-9611-ac1034fb3f79.pdf"}],"financialInterests":"","formattedTitle":"Cytogenetic analysis of cryptic species in allopatric population (2n = 60, NF = 82) of Nannospalax sp. (Spalacidae, Rodentia): conventional karyotype, constitutive heterochromatin, and NOR localization","fulltext":[{"header":"Introduction","content":"\u003cp\u003eMole rats (family Spalacidae) are the major component of steppe habitats in the Palearctic region (Topachevski 1976; Savic and Soldatovic \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1977\u003c/span\u003e), and is the most speciation groups taxon, which is currently regarded to be composed of more than 60 chromosomal forms in 2 genera (\u003cem\u003eSpalax\u003c/em\u003e and \u003cem\u003eNannospalax\u003c/em\u003e) in distributon area and the genus \u003cem\u003eNannospalax\u003c/em\u003e includes three morphospecies: \u003cem\u003eNannospalax leucodon\u003c/em\u003e, \u003cem\u003eNannospalax nehringi\u003c/em\u003e, and \u003cem\u003eNannospalax ehrenbergi\u003c/em\u003e in Turkey (Ellerman and Morrison-Scoot 1951; Topachevski 1976; Musser and Carleton \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Despite almost 30\u0026ndash;40\u0026nbsp;million years of the family Spalacidae evolutionary background, speciation is still in progress and many populations are at different levels of the process (Savic and Nevo \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Nevo et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; \u0026Uuml;nay \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Şen and Sarica 2011). The genus \u003cem\u003eNannospalax\u003c/em\u003e which is one of the youngest genera (about 24\u0026nbsp;million years ago) of this family (Hadid et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), shows a relatively high rate of karyotypic rearrangements and molecular divergence among mammals. These evolutionary dynamics provoke taxonomic uncertainties on the one hand but make this group a perfect model for evolutionary research on the other hand (Savic and Nevo \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Nevo et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; He et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Overall, the \u003cem\u003eNannospalax\u003c/em\u003e mole rats abound with cryptic species (Nevo et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Csorba et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Savic et al. 2017; Bugarski-Stanojević at al. 2022, 2024). Cryptic species are those that cannot readily be distinguished on the basis of phenotypic variation alone (Struck et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2018\u003c/span\u003e); they have created challenges in myriad areas of the life sciences, especially in taxonomic systems. Not all \u0026ldquo;cryptic\u0026rdquo; species, however, are truly cryptic; many are simply underexplored morphologically (Nastasi et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Cryptic species complexes where there is a large diversity of karyomorphs with autapomorphies must have originated due to plasticity in the ancestral karyomorph. The multiple karyotypic forms found in cryptic species could constitute, therefore, important examples of non-adaptive or adaptive chromosomal radiation, depending on the relation of the character with the environment (Kavalco and Pasa \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eChromosomal studies have often significantly contributed to understanding of the phylogeny and systematics of small mammals owing to extensive karyotypic variation recorded in their populations (Zima \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Arslan et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2011a\u003c/span\u003e). Changes in C-heterochromatin amount and distribution are among important mechanisms producing this variation. C- and Ag-NOR banding techniques are frequently used in animals and thus are useful for examining intra- and interspecific chromosomal differences between closely related species (Arslan et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2011a\u003c/span\u003e; Zhu et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The structure, number, and morphology of the nucleolar organizer region may be specific to populations, species, and subspecies. Nucleolus Organizer Regions (NORs) are frequently used as markers to show intra- and inter-species chromosomal polymorphism in animals. Species with limited gene exchange owing to geographic isolation have high NORs and chromosomal diversity. Therefore, even in small but isolated populations of these species, different karyotypes are found. The homogeneity of this character and the degree of diversity within a taxon largely determines the use of NORs in explaining relationships (Y\u0026uuml;ksel and Gaffaroğlu \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2008\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eConventional karyological studies carried out in Malatya province: Y\u0026uuml;ksel (\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e1984\u003c/span\u003e) 2n\u0026thinsp;=\u0026thinsp;60, NF\u0026thinsp;=\u0026thinsp;80 from Yazıhan; G\u0026uuml;lka\u0026ccedil; and Y\u0026uuml;ksel (\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1989\u003c/span\u003e) 2n\u0026thinsp;=\u0026thinsp;60, NF\u0026thinsp;=\u0026thinsp;80 from Ak\u0026ccedil;adağ and NF\u0026thinsp;=\u0026thinsp;82 from Arguvan; Nevo et al. (\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e1995\u003c/span\u003e) 2n\u0026thinsp;=\u0026thinsp;60, NF\u0026thinsp;=\u0026thinsp;78 from 30 km west of Malatya; Ivanitskaya et al. (\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1997\u003c/span\u003e) 2n\u0026thinsp;=\u0026thinsp;60, NF\u0026thinsp;=\u0026thinsp;78 from 12 km east of Malatya; Ulut\u0026uuml;rk et al. (\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) 2n\u0026thinsp;=\u0026thinsp;60, NF\u0026thinsp;=\u0026thinsp;82 from Arguvan, Hekimhan, and Arapgir; Coşkun et al. (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) 2n\u0026thinsp;=\u0026thinsp;60, NF\u0026thinsp;=\u0026thinsp;78 from Doğanşehir, Kale and Ak\u0026ccedil;adağ (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSo far C-, and NOR banding of different chromosomal forms of \u003cem\u003eNannospalax\u003c/em\u003e 2n\u0026thinsp;=\u0026thinsp;60 were achieved from Turkey by various authors (Ivanitskaya et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1997\u003c/span\u003e, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; G\u0026uuml;lka\u0026ccedil; and K\u0026uuml;\u0026ccedil;\u0026uuml;kdumlu \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Arslan and B\u0026ouml;l\u0026uuml;kbaş \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Arslan et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2011b\u003c/span\u003e; Aşan Baydemir et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Yağcı \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). However, there is not any banded karyotypes of \u003cem\u003eNannospalax\u003c/em\u003e population (2n\u0026thinsp;=\u0026thinsp;60, NF\u0026thinsp;=\u0026thinsp;82) in Arapgir (Malatya) region. The aim of the present study is to describe in detail the conventional chromosome staining and the distribution of the NORs and C-heterochromatin in the karyotype of \u003cem\u003eNannospalax sp.\u003c/em\u003e from Arapgir (Malatya).\u003c/p\u003e"},{"header":"Material and methods","content":"\u003cp\u003eCytogenetic analyses were performed on three specimens of mole rat from Arapgir, Malatya (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Karyotype preparations were obtained from the bone marrow of animals treated with colchicine (Ford and Hamerton \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1956\u003c/span\u003e). After preparation of chromosome slides, air-dried preparations were stained conventionally with Giemsa. Constitutive heterochromatin and nucleolus organizer regions were detected in individual autosomal and sex chromosome pairs via C-banding (Sumner, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e1972\u003c/span\u003e), and Ag-NOR staining (Howell and Black \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1980\u003c/span\u003e). From each specimen slides were prepared, and well-spread metaphase plates were analysed. The results were compared with previously published accounts on 2n\u0026thinsp;=\u0026thinsp;60 chromosomal forms carried out in Turkey. Standard voucher specimens (skins and skulls) are deposited in the Department of Biology, Faculty of Science, Dicle University, Diyarbakır, and Batman University Science and Art Faculty, Biology Department, Turkey.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003eConventional karyotype\u003c/h2\u003e \u003cp\u003eAll karyotyped mole rats from the population of Arapgir had identical karyotypes with 2n\u0026thinsp;=\u0026thinsp;60, NF\u0026thinsp;=\u0026thinsp;82 and NFa\u0026thinsp;=\u0026thinsp;78. The autosomal set consisted of ten pairs submetacentric (chromosome nos. 1\u0026ndash;10) and 19 pairs large-to-small acrocentric pairs (chromosome nos.11\u0026ndash;29) of gradually decreasing size). The X chromosome was the second large submetacentric, and the Y chromosome was dot-sized acrocentric (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eNOR banding patterns\u003c/h3\u003e\n\u003cp\u003eNORs were found in two bi-armed autosomal pairs (pairs 2 and 7) in the complements of all the specimens, and in the telomeric region of the short arms of those pairs, and only one homologue of the pairs numbers 4 and 10 autosomes has determined a positive silver signal (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e\n\u003ch3\u003eC-banding\u003c/h3\u003e\n\u003cp\u003eIn mole rats of Arapgir, C-positive heterochromatic blocks were located in centromeric region of all biarmed autosomes (1\u0026ndash;10) and while the one of the X chromosomes has a large pericentromeric C-positive region, the other one X chromosome has a dot like centromeric C positive (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e)\u003c/p\u003e \u003cp\u003eThe distribution of C-heterochromatin followed the same pattern in all examined specimen sets. Some biarmed autosome pairs (numbers 1, 5, 7, and 8) exhibited distinct dark centromeric C-bands. Other biarmed autosome pairs (numbers 2, 3, 4, 6, 9, and 10) exhibited a weaker C-positive band.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe 2n\u0026thinsp;=\u0026thinsp;60 chromosomal form shows the largest distribution areas among chromosomal forms of \u003cem\u003eNannospalax\u003c/em\u003e in Anatolia. The 2n\u0026thinsp;=\u0026thinsp;60 chromosomal form in Turkey were first recorded by Y\u0026uuml;ksel (\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e1984\u003c/span\u003e) from Malatya Province in the central-eastern Anatolia. Later, 2n\u0026thinsp;=\u0026thinsp;60, the most common chromosomal form of mole rats in central Anatolia, had been reported by numerous authors (G\u0026uuml;lka\u0026ccedil; and Y\u0026uuml;ksel \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1989\u003c/span\u003e; Nevo et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Ivanitskaya et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; G\u0026uuml;lka\u0026ccedil; and K\u0026uuml;\u0026ccedil;\u0026uuml;kdumlu \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Ulut\u0026uuml;rk et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Coşkun et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2010\u003c/span\u003e, and see Table \u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The karyotype in the 2n\u0026thinsp;=\u0026thinsp;60 chromosomal forms are variable and NF range of 72\u0026ndash;84 was reported by previous authors (Matur and S\u0026ouml;zen \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Kankılı\u0026ccedil; et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2007\u003c/span\u003e, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2009\u003c/span\u003e, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; S\u0026ouml;zen et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2011\u003c/span\u003e, \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Arslan et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). However, the geographical continuity of the known populations is in doubt. In the Malatya province two different chromosomal forms of 2n\u0026thinsp;=\u0026thinsp;60 (2n\u0026thinsp;=\u0026thinsp;60a, NF\u0026thinsp;=\u0026thinsp;82 and 2n\u0026thinsp;=\u0026thinsp;60b, NF\u0026thinsp;=\u0026thinsp;78) reported by Coşkun et al. (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), and they claimed that these two forms seem to be separated by the Tohma Stream.\u003c/p\u003e \u003cp\u003eThe NORs of \u003cem\u003eNannospalax\u003c/em\u003e populations in Malatya province were first studied by Ivanitskaya et al. (\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). According to these researchers, in all the samples, the NORs were localised in the short arms of 3 pairs of sub-telocentric chromosomes in this population. G\u0026uuml;lka\u0026ccedil; and K\u0026uuml;\u0026ccedil;\u0026uuml;kdumlu (\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1999\u003c/span\u003e) determined that \u003cem\u003eNannospalax\u003c/em\u003e (\u003cem\u003eSpalax leucodon\u003c/em\u003e) specimens obtained from Malatya, the karyotype consitsts of 30 chromosome pairs and the NORs were terminally located on 3 pairs of large subtelocentrics. All of the individuals displayed a consistent number of NORs per cell. NORs were not found on the sex chromosomes of \u003cem\u003eNannospalax\u003c/em\u003e (\u003cem\u003eSpalax leucodon\u003c/em\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eKaryotype records of the 2n\u0026thinsp;=\u0026thinsp;60 chromosomal forms of \u003cem\u003eNannospalax\u003c/em\u003e from Turkey. NF \u0026ndash; fundamental number of chromosomal arms (See Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e for location of the sites).\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"11\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003e2n\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNF\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003em/sm/st\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003ea\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eX\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eY\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eC band\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eNOR\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c10\" namest=\"c9\"\u003e \u003cp\u003eLocality\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c11\"\u003e \u003cp\u003eReference\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"22\" rowspan=\"23\"\u003e \u003cp\u003e\u003cb\u003e2n\u0026thinsp;=\u0026thinsp;60\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e80\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003esm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003est\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eMalatya\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eYazıhan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eY\u0026uuml;ksel \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e1984\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e80\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003esm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003est\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eMalatya\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eAk\u0026ccedil;adağ\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eG\u0026uuml;lka\u0026ccedil; and Y\u0026uuml;ksel \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1989\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e82\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003esm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ea\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eArguvan\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e78\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003esm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ea\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eMalatya\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e30 km W\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eNevo at al. 1995\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e78\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e14 m/sm\u0026thinsp;+\u0026thinsp;8a\u0026thinsp;+\u0026thinsp;XY\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e6 m/sm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eMalatya\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e12 km E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eIvanitskaya et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1997\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e6 m/sm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eMalatya\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eG\u0026uuml;lka\u0026ccedil; and K\u0026uuml;\u0026ccedil;\u0026uuml;kdumlu \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1999\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e78\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eSm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003ea\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e16\u0026thinsp;+\u0026thinsp;XY\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e8 m/sm (W)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u0026Ccedil;ankırı,\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eOrta, Sanabozu\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eIvanitskaya et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2008\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e9 m/sm (R)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eBursa\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eİneg\u0026ouml;l\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"3\" rowspan=\"4\"\u003e \u003cp\u003e82\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"3\" rowspan=\"4\"\u003e \u003cp\u003e20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\" morerows=\"3\" rowspan=\"4\"\u003e \u003cp\u003e38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ea\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eMalatya\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eArguvan, Hekimhan, Arapgir\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eUlut\u0026uuml;rk et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2009\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c5\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eSm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003ea\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eElazığ,\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eKeban (Denizli)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\" morerows=\"4\" rowspan=\"5\"\u003e \u003cp\u003eCoşkun et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2010\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eErzincan,\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e\u0026Ccedil;itk\u0026ouml;y, Dutluca\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eSivas\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eDivriği\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e78\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003esm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003ea\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eElazığ,\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eKeban (\u0026Ccedil;irkan)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eMalatya\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eDoğanşehir, Kale, Ak\u0026ccedil;adağ\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e76\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e44\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003esm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e14 m/sm\u0026thinsp;+\u0026thinsp;XY\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e8 m/sm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eAksaray\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eG\u0026uuml;lağa\u0026ccedil;, Ortak\u0026ouml;y, Ağa\u0026ccedil;\u0026ouml;ren\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eArslan and B\u0026ouml;l\u0026uuml;kbaş \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2010\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e78\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003esm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003em\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e16 m/sm\u0026thinsp;+\u0026thinsp;XY\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eG\u0026uuml;zelyurt, Sarıyahşi, Merkez\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e74\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e46\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003est\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e12 m/sm\u0026thinsp;+\u0026thinsp;XY\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e8 m/sm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\" morerows=\"3\" rowspan=\"4\"\u003e \u003cp\u003eKonya\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eHadim, Karatay\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\" morerows=\"3\" rowspan=\"4\"\u003e \u003cp\u003eArslan et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2011b\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e78\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e16m/sm\u0026thinsp;+\u0026thinsp;XY\u0026thinsp;+\u0026thinsp;10a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e8 m/sm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eIlgın, H\u0026ouml;y\u0026uuml;k, Saray\u0026ouml;n\u0026uuml;\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e79\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e21\u0026ndash;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003est\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e17\u0026thinsp;+\u0026thinsp;XY\u0026thinsp;+\u0026thinsp;12a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e8 m/sm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eBozkır, \u0026Ccedil;umra, Sel\u0026ccedil;uklu, G\u0026uuml;neysınır, Meram\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e80\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSt\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e18\u0026thinsp;+\u0026thinsp;XY\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e8 m/sm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eCihanbeyli\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e80\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ea\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e8 m/sm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eKonya\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eCihanbeyli, Kulu\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eAşan Baydemir et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2013\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e--\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e18 m/sm\u0026thinsp;+\u0026thinsp;XY\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e6 m/sm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eBilecik\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eS\u0026ouml;ğ\u0026uuml;t, Pazaryeri\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eYağcı \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2017\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e82\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ea\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e20 m/sm\u0026thinsp;+\u0026thinsp;XX\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e6 m/sm\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eMalatya\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eArapgir\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e\u003cb\u003eThis study\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eNevo et al. (\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e1995\u003c/span\u003e)d lka\u0026ccedil; and K\u0026uuml;\u0026ccedil;\u0026uuml;kdumlu (1999) reported that the NORs on homologue chromosomes do not exhibit heteromorphism in \u003cem\u003eNannospalax\u003c/em\u003e (\u003cem\u003eSpalax leucodon\u003c/em\u003e). The number of NORs carrying chromosomes and the location of the NORs on the chromosomes usually show remarkable variation among species. S\u0026ouml;zen et al. (\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2006\u003c/span\u003e) reported that it appears that changes in NORs have occurred in strict association with differentiation and evolution of species. Ivanitskaya et al. (\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2008\u003c/span\u003e) reported that both 2n\u0026thinsp;=\u0026thinsp;60R and 2n\u0026thinsp;=\u0026thinsp;60W chromosomal forms have NORs localized in the five pairs of sub-telocentric autosomes. Arslan et al. (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2011b\u003c/span\u003e) reported the NORs have in 4 pair autosomes of the four chromosomal forms (NF\u0026thinsp;=\u0026thinsp;74, 78, 79 and 80) of 2n\u0026thinsp;=\u0026thinsp;60 from Konya\u003c/p\u003e \u003cp\u003eDistributions of NORs patterns were very similar to those in Malatya (Ivanitskaya et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; G\u0026uuml;lka\u0026ccedil; and K\u0026uuml;\u0026ccedil;\u0026uuml;kdumlu \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1999\u003c/span\u003e) and Bilecik (Yağcı \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), but some differences in Bursa, \u0026Ccedil;ankırı (Ivanitskaya et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), Aksaray (Arslan and B\u0026ouml;l\u0026uuml;kbaş \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), Konya (Arslan et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2011b\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cem\u003eNannospalax\u003c/em\u003e (\u003cem\u003eS. leucodon\u003c/em\u003e) samples (2n\u0026thinsp;=\u0026thinsp;60, NF\u0026thinsp;=\u0026thinsp;78) from Malatya, all subtelocentric chromosomes possess completely heterochromatic short arms, with the exception of the pair 7; among the acrocentric chromosomes only four pair; have pericentromeric blocks of heterochromatin; in the submetacentric X-chromosomes the pericentromeric blocks of heterochromatin are not as dark as in the autosomes (Ivanitskaya et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). Chromosomes of blind mole rats from Malatya (2n\u0026thinsp;=\u0026thinsp;60, NF\u0026thinsp;=\u0026thinsp;78, NFa\u0026thinsp;=\u0026thinsp;74) given by Ivanitskaya et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1997\u003c/span\u003e differed distinctly with the C-banding pattern of Arapgir population (2n\u0026thinsp;=\u0026thinsp;60, NF\u0026thinsp;=\u0026thinsp;82, NFa\u0026thinsp;=\u0026thinsp;78).\u003c/p\u003e \u003cp\u003eIn terms of biarmed chromosomes, C-heterochromatin distributions were very similar those in \u0026Ccedil;ankırı and Bursa (Ivanitskaya et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), Aksaray (Arslan and B\u0026ouml;l\u0026uuml;kbaş \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) and Konya (Arslan et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2011b\u003c/span\u003e), but some differences in Malatya (Ivanitskaya et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). According to the acrocentric chromosomes C-heterochromatin distributions differ from in Malatya (Ivanitskaya et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1997\u003c/span\u003e) and Konya (Ilgın, H\u0026ouml;y\u0026uuml;k, Saray\u0026ouml;n\u0026uuml;, Bozkır, \u0026Ccedil;umra, Sel\u0026ccedil;uklu, G\u0026uuml;neysınır and Meram) (Arslan et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2011b\u003c/span\u003e). All sex chromosomes possesses C-heterochromatin blocks in all 2n\u0026thinsp;=\u0026thinsp;60 populations in Anatolia (See Table \u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Nucleolus organizer regions and C-heterochromatin distributions have been detected for the first time in this study for \u003cem\u003eNannospalax sp.\u003c/em\u003e 2n\u0026thinsp;=\u0026thinsp;60, NF\u0026thinsp;=\u0026thinsp;82 chromosomal form of Turkey.\u003c/p\u003e \u003cp\u003eTo date, no detailed cytogenetic investigations have been performed on representatives of \u003cem\u003eNannospalax sp.\u003c/em\u003e which inhabit a small territory in the northern part of the Tohma River basin in Arapgir (Malatya). Coşkun et al (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) stated that the two cryptic species (2n\u0026thinsp;=\u0026thinsp;60a, NF\u0026thinsp;=\u0026thinsp;82 and 2n\u0026thinsp;=\u0026thinsp;60b, NF\u0026thinsp;=\u0026thinsp;78) of mole rats have distributed in Malatya province and they argued the Tohma River acts as a barrier between two cryptic chromosomal forms. In this study, the presence of chromosomal forms 2n\u0026thinsp;=\u0026thinsp;60a, NF\u0026thinsp;=\u0026thinsp;82 (\u003cem\u003eNannospalax sp.\u003c/em\u003e) was confirmed. Morphological examination of specimens from both chromosomal forms is necessary to reveal morphological differences and, at last to a formal species description on this cryptic species. Our study demonstrates the uncovering hidden biodiversity. We, therefore, classified these mole rats as \u003cem\u003eNannospalax sp.\u003c/em\u003e but increasing the sample size is needed to rule out of this cryptic species.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eFunding\u003c/h2\u003e \u003cp\u003eThe authors declare that no funds, grants, or other support was received during the preparation of this manuscript.\u003c/p\u003e \u003cp\u003e \u003cb\u003eConflict of interest\u003c/b\u003e The authors declare no conflict of interest.\u003c/p\u003e \u003cp\u003e \u003cb\u003eEthical approval\u003c/b\u003e No ethical approval is required, not applicable.\u003c/p\u003e\u003ch2\u003eAuthor contribution\u003c/h2\u003e \u003cp\u003eAll authors made equal contributions to this study.\u003c/p\u003e\u003ch2\u003eData availability\u003c/h2\u003e \u003cp\u003eAll data supporting the findings of this study are available within the paper.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eArslan A, B\u0026ouml;l\u0026uuml;kbaş F (2010) C-heterochromatin and NORs distrubution of mole rat, \u003cem\u003eNannospalax xanthodon\u003c/em\u003e from Aksaray. 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Hystrix It J Mamm 11:5\u0026ndash;15. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.4404/hystrix-11.2-4143\u003c/span\u003e\u003cspan address=\"10.4404/hystrix-11.2-4143\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"biologia","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"biol","sideBox":"Learn more about [Biologia](http://link.springer.com/journal/11756)","snPcode":"11756","submissionUrl":"https://www.editorialmanager.com/biol/default2.aspx","title":"Biologia","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Nannospalax, mole rat, karyotype, Malatya, Turkey","lastPublishedDoi":"10.21203/rs.3.rs-6277258/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6277258/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eIn the present study, the 2n\u0026thinsp;=\u0026thinsp;60, NF\u0026thinsp;=\u0026thinsp;82 chromosomal form of blind mole rats, \u003cem\u003eNannospalax sp.\u003c/em\u003e, from Arapgir-Malatya Province in Turkey was investigated. Conventional chromosome staining, NOR (Nucleolus Organizer Region), and C-banding analysis were carried out on specimens of this population. The karyotype including 10 submetacentric pairs and 19 acrocentric pairs of autosomes and fundamental number is (NFa) 78. In all of the specimens studied the NOR\u0026rsquo;s were localized in the telomeric regions of the short arms of the 2 autosome pairs (nos. 2 and 7), and only one homologue of the pairs nos. 4 and 10 autosomes has a positive silver signal. C-heterochromatin regions were found in the centromeric region of whole bi-armed autosomal pairs. One of the X chromosomes has a pericentromeric C-positive band, whereas the other has a dot-like C-positive band.\u003c/p\u003e","manuscriptTitle":"Cytogenetic analysis of cryptic species in allopatric population (2n = 60, NF = 82) of Nannospalax sp. (Spalacidae, Rodentia): conventional karyotype, constitutive heterochromatin, and NOR localization","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-06-26 14:03:38","doi":"10.21203/rs.3.rs-6277258/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Major revisions","date":"2025-07-26T08:14:16+00:00","index":"","fulltext":""},{"type":"reviewerAgreed","content":"","date":"2025-06-24T11:39:15+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-06-24T09:29:36+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Biologia","date":"2025-06-08T13:53:14+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-03-25T11:49:11+00:00","index":"","fulltext":""},{"type":"submitted","content":"Biologia","date":"2025-03-21T15:10:16+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"biologia","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"biol","sideBox":"Learn more about [Biologia](http://link.springer.com/journal/11756)","snPcode":"11756","submissionUrl":"https://www.editorialmanager.com/biol/default2.aspx","title":"Biologia","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"dd5577a7-1390-4820-9809-b2d7e3e52ca9","owner":[],"postedDate":"June 26th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2026-01-12T16:01:04+00:00","versionOfRecord":{"articleIdentity":"rs-6277258","link":"https://doi.org/10.1007/s11756-025-02124-x","journal":{"identity":"biologia","isVorOnly":false,"title":"Biologia"},"publishedOn":"2026-01-08 15:57:22","publishedOnDateReadable":"January 8th, 2026"},"versionCreatedAt":"2025-06-26 14:03:38","video":"","vorDoi":"10.1007/s11756-025-02124-x","vorDoiUrl":"https://doi.org/10.1007/s11756-025-02124-x","workflowStages":[]},"version":"v1","identity":"rs-6277258","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6277258","identity":"rs-6277258","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}