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1

Freedberg, Steven, Rachel M. Bowden, Michael A. Ewert, Dale R. Sengelaub, and Craig E. Nelson. "Long-term sex reversal by oestradiol in amniotes with heteromorphic sex chromosomes." Biology Letters 2, no. 3 (2006): 378–81. http://dx.doi.org/10.1098/rsbl.2006.0454.

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Oestradiol application during embryonic development reverses the sex of male embryos and results in normal female differentiation in reptiles lacking heteromorphic sex chromosomes, but fails to do so in birds and mammals with heteromorphic sex chromosomes. It is not clear whether the evolution of heteromorphic sex chromosomes in amniotes is accompanied by insensitivity to oestradiol, or if the association between oestradiol insensitivity and heteromorphic sex chromosomes can be attributable to phylogenetic constraints in these taxa. Turtles provide an ideal system to examine the potential rela
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2

Ogata, Mitsuaki, Kazuo Suzuki, Yoshiaki Yuasa, and Ikuo Miura. "Sex chromosome evolution from a heteromorphic to a homomorphic system by inter-population hybridization in a frog." Philosophical Transactions of the Royal Society B: Biological Sciences 376, no. 1833 (2021): 20200105. http://dx.doi.org/10.1098/rstb.2020.0105.

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Sex chromosomes generally evolve from a homomorphic to heteromorphic state. Once a heteromorphic system is established, the sex chromosome system may remain stable for an extended period. Here, we show the opposite case of sex chromosome evolution from a heteromorphic to a homomorphic system in the Japanese frog Glandirana rugosa. One geographic group, Neo-ZW, has ZZ-ZW type heteromorphic sex chromosomes. We found that its western edge populations, which are geographically close to another West-Japan group with homomorphic sex chromosomes of XX-XY type, showed homozygous genotypes of sex-linke
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3

Ma, Wen-Juan, and Paris Veltsos. "The Diversity and Evolution of Sex Chromosomes in Frogs." Genes 12, no. 4 (2021): 483. http://dx.doi.org/10.3390/genes12040483.

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Frogs are ideal organisms for studying sex chromosome evolution because of their diversity in sex chromosome differentiation and sex-determination systems. We review 222 anuran frogs, spanning ~220 Myr of divergence, with characterized sex chromosomes, and discuss their evolution, phylogenetic distribution and transitions between homomorphic and heteromorphic states, as well as between sex-determination systems. Most (~75%) anurans have homomorphic sex chromosomes, with XY systems being three times more common than ZW systems. Most remaining anurans (~25%) have heteromorphic sex chromosomes, w
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4

Nielsen, Stuart V., Brendan J. Pinto, Irán Andira Guzmán-Méndez, and Tony Gamble. "First Report of Sex Chromosomes in Night Lizards (Scincoidea: Xantusiidae)." Journal of Heredity 111, no. 3 (2020): 307–13. http://dx.doi.org/10.1093/jhered/esaa007.

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Abstract Squamate reptiles (lizards, snakes, and amphibians) are an outstanding group for studying sex chromosome evolution—they are old, speciose, geographically widespread, and exhibit myriad sex-determining modes. Yet, the vast majority of squamate species lack heteromorphic sex chromosomes. Cataloging the sex chromosome systems of species lacking easily identifiable, heteromorphic sex chromosomes, therefore, is essential before we are to fully understand the evolution of vertebrate sex chromosomes. Here, we use restriction site-associated DNA sequencing (RADseq) to classify the sex chromos
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5

Stevens, Lewis, Manuela Kieninger, Brian Chan, et al. "The genome of Litomosoides sigmodontis illuminates the origins of Y chromosomes in filarial nematodes." PLOS Genetics 20, no. 1 (2024): e1011116. http://dx.doi.org/10.1371/journal.pgen.1011116.

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Heteromorphic sex chromosomes are usually thought to have originated from a pair of autosomes that acquired a sex-determining locus and subsequently stopped recombining, leading to degeneration of the sex-limited chromosome. The majority of nematodes species lack heteromorphic sex chromosomes and determine sex using an X-chromosome counting mechanism, with males being hemizygous for one or more X chromosomes (XX/X0). Some filarial nematode species, including important parasites of humans, have heteromorphic XX/XY karyotypes. It has been assumed that sex is determined by a Y-linked locus in the
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6

Vacarizas, Joshua, Takahiro Taguchi, Takuma Mezaki, Sam Edward Manalili, Rei Kawakami, and Satoshi Kubota. "Cytogenetic evidence and dmrt linkage indicate male heterogamety in a non-bilaterian animal." PLOS ONE 18, no. 5 (2023): e0285851. http://dx.doi.org/10.1371/journal.pone.0285851.

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The diversity of sex determination systems in animals suggests that sex chromosomes evolve independently across different lineages. However, the present data on these systems is largely limited and represented mainly by bilaterian animals. Sex chromosomes and sex determination system based on cytogenetic evidence remain a mystery among non-bilaterians, the most basal animals. Here, we investigated the sex determination system of a non-bilaterian (Goniopora djiboutiensis) based on karyotypic analysis and identification of locus of dmrt1, a known master sex-determining gene in many animals. Resu
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7

Darolti, Iulia, Alison E. Wright, Benjamin A. Sandkam, et al. "Extreme heterogeneity in sex chromosome differentiation and dosage compensation in livebearers." Proceedings of the National Academy of Sciences 116, no. 38 (2019): 19031–36. http://dx.doi.org/10.1073/pnas.1905298116.

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Once recombination is halted between the X and Y chromosomes, sex chromosomes begin to differentiate and transition to heteromorphism. While there is a remarkable variation across clades in the degree of sex chromosome divergence, far less is known about the variation in sex chromosome differentiation within clades. Here, we combined whole-genome and transcriptome sequencing data to characterize the structure and conservation of sex chromosome systems across Poeciliidae, the livebearing clade that includes guppies. We found that the Poecilia reticulata XY system is much older than previously t
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8

Crepaldi, Carolina, and Patricia P. Parise-Maltempi. "Heteromorphic Sex Chromosomes and Their DNA Content in Fish: An Insight through Satellite DNA Accumulation in Megaleporinus elongatus." Cytogenetic and Genome Research 160, no. 1 (2020): 38–46. http://dx.doi.org/10.1159/000506265.

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The repetitive DNA content of fish sex chromosomes provides valuable insights into specificities and patterns of their genetic sex determination systems. In this study, we revealed the genomic satellite DNA (satDNA) content of Megaleporinuselongatus, a Neotropical fish species with Z1Z1Z2Z2/Z1W1Z2W2 multiple sex chromosomes, through high-throughput analysis and graph-based clustering, isolating 68 satDNA families. By physically mapping these sequences in female metaphases, we discovered 15 of the most abundant satDNAs clustered in its chromosomes, 9 of which were found exclusively in the highl
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9

Mezzasalma, Marcello, Fabio M. Guarino, and Gaetano Odierna. "Lizards as Model Organisms of Sex Chromosome Evolution: What We Really Know from a Systematic Distribution of Available Data?" Genes 12, no. 9 (2021): 1341. http://dx.doi.org/10.3390/genes12091341.

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Lizards represent unique model organisms in the study of sex determination and sex chromosome evolution. Among tetrapods, they are characterized by an unparalleled diversity of sex determination systems, including temperature-dependent sex determination (TSD) and genetic sex determination (GSD) under either male or female heterogamety. Sex chromosome systems are also extremely variable in lizards. They include simple (XY and ZW) and multiple (X1X2Y and Z1Z2W) sex chromosome systems and encompass all the different hypothesized stages of diversification of heterogametic chromosomes, from homomor
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10

Luykx, Peter. "Variation in sex-linked interchange heterozygosity in the termite Incisitermes schwarzi Banks (Insecta: Isoptera) on the island of Jamaica." Genome 29, no. 2 (1987): 319–25. http://dx.doi.org/10.1139/g87-052.

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Collections of colonies of the termite Incisitermes schwarzi from mangroves around the coast of Jamaica revealed six chromosomal types, all involving variations or rearrangements of the sex chromosomes. One of the types had a heteromorphic sex bivalent in which the Y chromosome was larger than the X. The other five races had complex interchange multiples: a chain of 11, a chain of 12, a ring of 12, a ring of 14, and a ring of 18 chromosomes. The situation is similar to that described previously for Kalotermes approximatus, another member of the family Kalotermitidae, in the southeastern United
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11

Ogata, Mitsuaki, Foyez Shams, Yuri Yoshimura, Tariq Ezaz, and Ikuo Miura. "W Chromosome Evolution by Repeated Recycling in the Frog Glandirana rugosa." DNA 2, no. 3 (2022): 172–84. http://dx.doi.org/10.3390/dna2030012.

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The Y or W sex chromosome of a heteromorphic pair is usually heterochromatinised and degenerated. However, whether chromosome degeneration constantly proceeds toward an extreme end is not fully understood. Here, we present a case of intermittent evolution of W chromosomes caused by interpopulation hybridisation in the Japanese soil-frog, Glandirana rugosa. This species includes two heteromorphic sex chromosome systems, which are separated into geographic populations, namely the XY and ZW groups. In this study, to uncover the evolutionary mechanisms of the heterogeneous W chromosomes, we geneti
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12

Bista, Basanta, and Nicole Valenzuela. "Turtle Insights into the Evolution of the Reptilian Karyotype and the Genomic Architecture of Sex Determination." Genes 11, no. 4 (2020): 416. http://dx.doi.org/10.3390/genes11040416.

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Sex chromosome evolution remains an evolutionary puzzle despite its importance in understanding sexual development and genome evolution. The seemingly random distribution of sex-determining systems in reptiles offers a unique opportunity to study sex chromosome evolution not afforded by mammals or birds. These reptilian systems derive from multiple transitions in sex determination, some independent, some convergent, that lead to the birth and death of sex chromosomes in various lineages. Here we focus on turtles, an emerging model group with growing genomic resources. We review karyotypic chan
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13

Salgado, Filipe Schitini, Marina Souza Cunha, Silvana Melo, and Jorge Abdala Dergam. "Cytogenetic analysis of Hypomasticus copelandii and H. steindachneri: relevance of cytotaxonomic markers in the Anostomidae family (Characiformes)." Comparative Cytogenetics 15, no. 1 (2021): 65–76. http://dx.doi.org/10.3897/compcytogen.v15.i1.61957.

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Recent phylogenetic hypotheses within Anostomidae, based on morphological and molecular data, resulted in the description of new genera (Megaleporinus Ramirez, Birindelli et Galetti, 2017) and the synonymization of others, such as the reallocation of Leporinus copelandii Steindachner, 1875 and Leporinus steindachneri Eigenmann, 1907 to Hypomasticus Borodin, 1929. Despite high levels of conservatism of the chromosomal macrostructure in this family, species groups have been corroborated using banding patterns and the presence of different sex chromosome systems. Due to the absence of cytogenetic
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14

Peralta, Diego M., Juan I. Túnez, Ulises E. Rodríguez Cruz, and Santiago G. Ceballos. "A rapid approach for sex assignment by RAD-seq using a reference genome." PLOS ONE 19, no. 4 (2024): e0297987. http://dx.doi.org/10.1371/journal.pone.0297987.

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Sex identification is a common objective in molecular ecology. While many vertebrates display sexual dimorphism, determining the sex can be challenging in certain situations, such as species lacking clear sex-related phenotypic characteristics or in studies using non-invasive methods. In these cases, DNA analyses serve as valuable tools not only for sex determination but also for validating sex assignment based on phenotypic traits. In this study, we developed a bioinformatic framework for sex assignment using genomic data obtained through GBS, and having an available closely related genome as
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15

Sassi, Francisco de M. C., Orlando Moreira-Filho, Geize A. Deon, et al. "Adding New Pieces to the Puzzle of Karyotype Evolution in Harttia (Siluriformes, Loricariidae): Investigation of Amazonian Species." Biology 10, no. 9 (2021): 922. http://dx.doi.org/10.3390/biology10090922.

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A remarkable morphological diversity and karyotype variability can be observed in the Neotropical armored catfish genus Harttia. These fishes offer a useful model to explore both the evolution of karyotypes and sex chromosomes, since many species possess male-heterogametic sex chromosome systems and a high rate of karyotype repatterning. Based on the karyotype organization, the chromosomal distribution of several repetitive DNA classes, and the rough estimates of genomic divergences at the intraspecific and interspecific levels via Comparative Genomic Hybridization, we identified shared diploi
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16

Salgado, Filipe Schitini, Marina Souza Cunha, Silvana Melo, and Jorge Abdala Dergam. "Cytogenetic analysis of Hypomasticus copelandii and H. steindachneri: relevance of cytotaxonomic markers in the Anostomidae family (Characiformes)." Comparative Cytogenetics 15, no. (1) (2021): 65–76. https://doi.org/10.3897/compcytogen.v15.i1.61957.

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Recent phylogenetic hypotheses within Anostomidae, based on morphological and molecular data, resulted in the description of new genera (Megaleporinus Ramirez, Birindelli et Galetti, 2017) and the synonymization of others, such as the reallocation of Leporinus copelandii Steindachner, 1875 and Leporinus steindachneri Eigenmann, 1907 to Hypomasticus Borodin, 1929. Despite high levels of conservatism of the chromosomal macrostructure in this family, species groups have been corroborated using banding patterns and the presence of different sex chromosome systems. Due to the absence of cytogenetic
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17

Mezzasalma, Marcello, Gaetano Odierna, Rachele Macirella, and Elvira Brunelli. "Comparative Cytogenetics of the Malagasy Ground Geckos of the Paroedura bastardi and Paroedura picta Species Groups." Animals 14, no. 11 (2024): 1708. http://dx.doi.org/10.3390/ani14111708.

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We present a comparative chromosome study of several taxa of the Malagasy ground geckos of the Paroedura bastardi and P. picta species groups. We employed a preliminary molecular analysis using a trait of the mitochondrial 16S rRNA gene (of about 570 bp) to assess the taxonomic status of the samples studied and a cytogenetic analysis with standard karyotyping (5% Giemsa solution), silver staining (Ag–NOR staining) and sequential C-banding (C-banding + Giemsa and + fluorochromes). Our results show that all the taxa studied of the P. bastardi group (P. ibityensis, P. rennerae and P. cf. guibeae)
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18

Panthum, Thitipong, Worapong Singchat, Nararat Laopichienpong, et al. "Genome-Wide SNP Analysis of Male and Female Rice Field Frogs, Hoplobatrachus rugulosus, Supports a Non-Genetic Sex Determination System." Diversity 13, no. 10 (2021): 501. http://dx.doi.org/10.3390/d13100501.

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Sex determination systems (SDSs) in anurans are diverse and have undergone independent evolutionary transitions among species. The mode of sexual reproduction of the rice field frog (Hoplobatrachus rugulosus)—an economically viable, edible amphibian species—is not well known. Previous studies have proposed that threshold temperature conditions may determine sex in these frogs. To elucidate the SDS in H. rugulosus, we karyotyped 10 male and 12 female frogs, and performed fluorescence in situ hybridization combined with sequencing analyses using DArTseq™. Our results revealed a highly conserved
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19

Souza, Fernando H. S. de, Francisco de M. C. Sassi, Pedro H. N. Ferreira, et al. "Integrating Cytogenetics and Population Genomics: Allopatry and Neo-Sex Chromosomes May Have Shaped the Genetic Divergence in the Erythrinus erythrinus Species Complex (Teleostei, Characiformes)." Biology 11, no. 2 (2022): 315. http://dx.doi.org/10.3390/biology11020315.

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Diversity found in Neotropical freshwater fish is remarkable. It can even hinder a proper delimitation of many species, with the wolf fish Erythrinus erythrinus (Teleostei, Characiformes) being a notable example. This nominal species shows remarkable intra-specific variation, with extensive karyotype diversity found among populations in terms of different diploid chromosome numbers (2n), karyotype compositions and sex chromosome systems. Here, we analyzed three distinct populations (one of them cytogenetically investigated for the first time) that differed in terms of their chromosomal feature
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20

Mawaribuchi, Shuuji, Michihiko Ito, Mitsuaki Ogata, Yuri Yoshimura, and Ikuo Miura. "Parallel Evolution of Sex-Linked Genes across XX/XY and ZZ/ZW Sex Chromosome Systems in the Frog Glandirana rugosa." Genes 14, no. 2 (2023): 257. http://dx.doi.org/10.3390/genes14020257.

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Genetic sex-determination features are male (XX/XY) or female heterogamety (ZZ/ZW). To identify similarities and differences in the molecular evolution of sex-linked genes between these systems, we directly compared the sex chromosome systems existing in the frog Glandirana rugosa. The heteromorphic X/Y and Z/W sex chromosomes were derived from chromosomes 7 (2n = 26). RNA-Seq, de novo assembly, and BLASTP analyses identified 766 sex-linked genes. These genes were classified into three different clusters (XW/YZ, XY/ZW, and XZ/YW) based on sequence identities between the chromosomes, probably r
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21

Waters, Paul D., and Jennifer A. Marshall Graves. "Monotreme sex chromosomes - implications for the evolution of amniote sex chromosomes." Reproduction, Fertility and Development 21, no. 8 (2009): 943. http://dx.doi.org/10.1071/rd09250.

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In vertebrates, a highly conserved pathway of genetic events controls male and female development, to the extent that many genes involved in human sex determination are also involved in fish sex determination. Surprisingly, the master switch to this pathway, which intuitively could be considered the most critical step, is inconsistent between vertebrate taxa. Interspersed in the vertebrate tree there are species that determine sex by environmental cues such as the temperature at which eggs are incubated, and then there are genetic sex-determination systems, with male heterogametic species (XY
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22

Santos da Silva, Kevin, Larissa Glugoski, Marcelo Ricardo Vicari, et al. "Mechanisms of Karyotypic Diversification in Ancistrus (Siluriformes, Loricariidae): Inferences from Repetitive Sequence Analysis." International Journal of Molecular Sciences 24, no. 18 (2023): 14159. http://dx.doi.org/10.3390/ijms241814159.

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Ancistrus is a highly diverse neotropical fish genus that exhibits extensive chromosomal variability, encompassing karyotypic morphology, diploid chromosome number (2n = 34–54), and the evolution of various types of sex chromosome systems. Robertsonian rearrangements related to unstable chromosomal sites are here described. Here, the karyotypes of two Ancistrus species were comparatively analyzed using classical cytogenetic techniques, in addition to isolation, cloning, sequencing, molecular characterization, and fluorescence in situ hybridization of repetitive sequences (i.e., 18S and 5S rDNA
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23

Rovatsos, Michail, Martina Johnson Pokorná, Marie Altmanová, and Lukáš Kratochvíl. "Mixed-Up Sex Chromosomes: Identification of Sex Chromosomes in the X1X1X2X2/X1X2Y System of the Legless Lizards of the Genus Lialis (Squamata: Gekkota: Pygopodidae)." Cytogenetic and Genome Research 149, no. 4 (2016): 282–89. http://dx.doi.org/10.1159/000450734.

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Geckos in general show extensive variability in sex determining systems, but only male heterogamety has been demonstrated in the members of their legless family Pygopodidae. In the pioneering study published more than 45 years ago, multiple sex chromosomes of the type X1X1X2X2/X1X2Y were described in Burton's legless lizard (Lialisburtonis) based on conventional cytogenetic techniques. We conducted cytogenetic analyses including comparative genomic hybridization and fluorescence in situ hybridization (FISH) with selected cytogenetic markers in this species and the previously cytogenetically un
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24

Furman, Benjamin L. S., David C. H. Metzger, Iulia Darolti, et al. "Sex Chromosome Evolution: So Many Exceptions to the Rules." Genome Biology and Evolution 12, no. 6 (2020): 750–63. http://dx.doi.org/10.1093/gbe/evaa081.

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Abstract Genomic analysis of many nonmodel species has uncovered an incredible diversity of sex chromosome systems, making it possible to empirically test the rich body of evolutionary theory that describes each stage of sex chromosome evolution. Classic theory predicts that sex chromosomes originate from a pair of homologous autosomes and recombination between them is suppressed via inversions to resolve sexual conflict. The resulting degradation of the Y chromosome gene content creates the need for dosage compensation in the heterogametic sex. Sex chromosome theory also implies a linear proc
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25

Romanenko, Svetlana A., Antonina V. Smorkatcheva, Yulia M. Kovalskaya, et al. "Complex Structure of Lasiopodomys mandarinus vinogradovi Sex Chromosomes, Sex Determination, and Intraspecific Autosomal Polymorphism." Genes 11, no. 4 (2020): 374. http://dx.doi.org/10.3390/genes11040374.

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The mandarin vole, Lasiopodomys mandarinus, is one of the most intriguing species among mammals with non-XX/XY sex chromosome system. It combines polymorphism in diploid chromosome numbers, variation in the morphology of autosomes, heteromorphism of X chromosomes, and several sex chromosome systems the origin of which remains unexplained. Here we elucidate the sex determination system in Lasiopodomys mandarinus vinogradovi using extensive karyotyping, crossbreeding experiments, molecular cytogenetic methods, and single chromosome DNA sequencing. Among 205 karyotyped voles, one male and three f
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Pajpach, Filip, Tianyu Wu, Linda Shearwin-Whyatt, Keith Jones, and Frank Grützner. "Flavors of Non-Random Meiotic Segregation of Autosomes and Sex Chromosomes." Genes 12, no. 9 (2021): 1338. http://dx.doi.org/10.3390/genes12091338.

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Segregation of chromosomes is a multistep process occurring both at mitosis and meiosis to ensure that daughter cells receive a complete set of genetic information. Critical components in the chromosome segregation include centromeres, kinetochores, components of sister chromatid and homologous chromosomes cohesion, microtubule organizing centres, and spindles. Based on the cytological work in the grasshopper Brachystola, it has been accepted for decades that segregation of homologs at meiosis is fundamentally random. This ensures that alleles on chromosomes have equal chance to be transmitted
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27

Radder, Rajkumar S., Alexander E. Quinn, Arthur Georges, Stephen D. Sarre, and Richard Shine. "Genetic evidence for co-occurrence of chromosomal and thermal sex-determining systems in a lizard." Biology Letters 4, no. 2 (2007): 176–78. http://dx.doi.org/10.1098/rsbl.2007.0583.

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An individual's sex depends upon its genes (genotypic sex determination or GSD) in birds and mammals, but reptiles are more complex: some species have GSD whereas in others, nest temperatures determine offspring sex (temperature-dependent sex determination). Previous studies suggested that montane scincid lizards ( Bassiana duperreyi , Scincidae) possess both of these systems simultaneously: offspring sex is determined by heteromorphic sex chromosomes (XX–XY system) in most natural nests, but sex ratio shifts suggest that temperatures override chromosomal sex in cool nests to generate phenotyp
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Lisachov, Artem P., Vladimir A. Trifonov, Massimo Giovannotti, Malcolm A. Ferguson-Smith, and Pavel M. Borodin. "Heteromorphism of “Homomorphic” Sex Chromosomes in Two Anole Species (Squamata, Dactyloidae) Revealed by Synaptonemal Complex Analysis." Cytogenetic and Genome Research 151, no. 2 (2017): 89–95. http://dx.doi.org/10.1159/000460829.

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Iguanians (Pleurodonta) are one of the reptile lineages that, like birds and mammals, have sex chromosomes of ancient origin. In most iguanians these are microchromosomes, making a distinction between the X and Y as well as between homeologous sex chromosomes in other species difficult. Meiotic chromosome analysis may be used to elucidate their differentiation, because meiotic prophase chromosomes are longer and less condensed than metaphase chromosomes, and the homologues are paired with each other, revealing minor heteromorphisms. Using electron and fluorescent microscopy of surface spread s
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Grant, Kirstie D., Daniel Koenemann, Janet Mansaray, et al. "A new phylogeny of Rumex (Polygonaceae) adds evolutionary context to the diversity of reproductive systems present in the genus." PhytoKeys 204 (August 5, 2022): 57–72. https://doi.org/10.3897/phytokeys.204.85256.

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Rumex is one of about 50 genera in the knotweed family, Polygonaceae. The genus comprises about 200 species with bisexual, or more commonly, unisexual flowers, with the species displaying monoecious, dioecious, synoecious (hermaphroditic) or polygamous reproductive systems. Some of the dioecious species have heteromorphic sex chromosomes, which is rare amongst angiosperms. We here present a plastid phylogeny of 67 species, representing all four subgenera. For this study, we used three chloroplast markers, rbcL, trnH-psbA, trnL-F and dense taxon sampling to reconstruct the most comprehensive mo
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Grant, Kirstie D., Daniel Koenemann, Janet Mansaray, et al. "A new phylogeny of Rumex (Polygonaceae) adds evolutionary context to the diversity of reproductive systems present in the genus." PhytoKeys 204 (August 5, 2022): 57–72. http://dx.doi.org/10.3897/phytokeys.204.85256.

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Rumex is one of about 50 genera in the knotweed family, Polygonaceae. The genus comprises about 200 species with bisexual, or more commonly, unisexual flowers, with the species displaying monoecious, dioecious, synoecious (hermaphroditic) or polygamous reproductive systems. Some of the dioecious species have heteromorphic sex chromosomes, which is rare amongst angiosperms. We here present a plastid phylogeny of 67 species, representing all four subgenera. For this study, we used three chloroplast markers, rbcL, trnH-psbA, trnL-F and dense taxon sampling to reconstruct the most comprehensive mo
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31

Chen, J., Y. Fu, D. Xiang, et al. "XX/XY heteromorphic sex chromosome systems in two bullhead catfish species, Liobagrusmarginatus and L. styani (Amblycipitidae, Siluriformes)." Cytogenetic and Genome Research 122, no. 2 (2008): 169–74. http://dx.doi.org/10.1159/000163095.

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32

Montiel, Eugenia Elisabet, Daleen Badenhorst, LingSze Lee, and Nicole Valenzuela. "Evolution and dosage compensation of nucleolar organizing regions (NORs) mediated by mobile elements in turtles with female (ZZ/ZW) but not with male (XX/XY) heterogamety." Journal of Evolutionary Biology 35, no. 12 (2022): 1709–20. https://doi.org/10.1111/jeb.14064.

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Understanding the evolution and regulation of nucleolar organizing regions (NORs) is important to elucidate genome structure and function. This is because ribosomal gene (rDNA) copy number and activity mediate protein biosynthesis, stress response, ageing, disease, dosage compensation and genome stability. Here, we found contrasting dosage compensation of sex-linked NORs in turtles with male and female heterogamety. Most taxa examined exhibit homomorphic rRNA gene clusters in a single autosome pair (determined by 28S rDNA fluorescence in situ hybridization), whereas NORs are sex-linked in <em>
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Yamamoto, Kayoko, Takashi Hamaji, Hiroko Kawai-Toyooka, et al. "Three genomes in the algal genus Volvox reveal the fate of a haploid sex-determining region after a transition to homothallism." Proceedings of the National Academy of Sciences 118, no. 21 (2021): e2100712118. http://dx.doi.org/10.1073/pnas.2100712118.

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Transitions between separate sexes (dioecy) and other mating systems are common across eukaryotes. Here, we study a change in a haploid dioecious green algal species with male- and female-determining chromosomes (U and V). The genus Volvox is an oogamous (with large, immotile female gametes and small, motile male gametes) and includes both heterothallic species (with distinct male and female genotypes, associated with a mating-type system that prevents fusion of gametes of the same sex) and homothallic species (bisexual, with the ability to self-fertilize). We date the origin of an expanded se
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34

Mahony, M. J. "Heteromorphic sex chromosomes in the Australian frog Crinia bilingua (Anura: Myobatrachidae)." Genome 34, no. 3 (1991): 334–37. http://dx.doi.org/10.1139/g91-055.

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The karyotype of Crinia bilingua was examined and analysed with standard staining, C-banding, and silver-staining. Heteromorphic sex chromosomes of the ZW ♂/ZZ ♀ type were observed. The larger W chromosome is submetacentric and the smaller Z chromosome is acrocentric. The centromere and proximal region of the short arm of the W chromosome consist of constitutive heterochromatin (C-band region), and beyond this is a small euchromatic terminal region. The centromere of the Z chromosome did not C-band. The long arms of the Z and W chromosomes are euchromatic and equal in length. The nucleolar org
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35

Sessions, Stanley K., Lilijana Bizjak Mali, David M. Green, Vladimir Trifonov, and Malcolm Ferguson-Smith. "Evidence for Sex Chromosome Turnover in Proteid Salamanders." Cytogenetic and Genome Research 148, no. 4 (2016): 305–13. http://dx.doi.org/10.1159/000446882.

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A major goal of genomic and reproductive biology is to understand the evolution of sex determination and sex chromosomes. Species of the 2 genera of the Salamander family Proteidae - Necturus of eastern North America, and Proteus of Southern Europe - have similar-looking karyotypes with the same chromosome number (2n = 38), which differentiates them from all other salamanders. However, Necturus possesses strongly heteromorphic X and Y sex chromosomes that Proteus lacks. Since the heteromorphic sex chromosomes of Necturus were detectable only with C-banding, we hypothesized that we could use C-
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36

Singchat, Worapong, Syed Farhan Ahmad, Nararat Laopichienpong, et al. "Snake W Sex Chromosome: The Shadow of Ancestral Amniote Super-Sex Chromosome." Cells 9, no. 11 (2020): 2386. http://dx.doi.org/10.3390/cells9112386.

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Heteromorphic sex chromosomes, particularly the ZZ/ZW sex chromosome system of birds and some reptiles, undergo evolutionary dynamics distinct from those of autosomes. The W sex chromosome is a unique karyological member of this heteromorphic pair, which has been extensively studied in snakes to explore the origin, evolution, and genetic diversity of amniote sex chromosomes. The snake W sex chromosome offers a fascinating model system to elucidate ancestral trajectories that have resulted in genetic divergence of amniote sex chromosomes. Although the principal mechanism driving evolution of th
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37

Kuwana, Chiao, Hiroyuki Fujita, Masataka Tagami, Takanori Matsuo, and Ikuo Miura. "Evolution of Sex Chromosome Heteromorphy in Geographic Populations of the Japanese Tago’s Brown Frog Complex." Cytogenetic and Genome Research 161, no. 1-2 (2021): 23–31. http://dx.doi.org/10.1159/000512964.

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The sex chromosomes of most anuran amphibians are characterized by homomorphy in both sexes, and evolution to heteromorphy rarely occurs at the species or geographic population level. Here, we report sex chromosome heteromorphy in geographic populations of the Japanese Tago’s brown frog complex (2&lt;i&gt;n&lt;/i&gt; = 26), comprising &lt;i&gt;Rana sakuraii&lt;/i&gt; and &lt;i&gt;R. tagoi&lt;/i&gt;. The sex chromosomes of &lt;i&gt;R. sakuraii&lt;/i&gt; from the populations in western Japan were homomorphic in both sexes, whereas chromosome 7 from the populations in eastern Japan were heteromor
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38

Thiriot-Quiévreux, Catherine, and Roger R. Seapy. "Chromosome studies of three families of pelagic heteropod molluscs (Atlantidae, Carinariidae, and Pterotracheidae) from Hawaiian waters." Canadian Journal of Zoology 75, no. 2 (1997): 237–44. http://dx.doi.org/10.1139/z97-030.

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Chromosome number and morphology were studied in gonadal tissue of 11 species of Atlantidae, 2 species of Carinariidae, and 3 species of Pterotracheidae, using an air-drying technique and Giemsa staining. In the Atlantidae the diploid chromosome number was the same in males and females and there were no heteromorphic chromosomes. The diploid chromosome number in nine species of Atlanta was 30 and the majority of chromosome pairs were metacentric and submetacentric. In Protatlanta souleyeti the diploid number was 28, and included five metacentric, six submetacentric, and three subtelocentric ch
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39

Mahesh, G., N. B. Ramachandra, and H. A. Ranganath. "Autoradiographic study of transcription and dosage compensation in the sex and neo-sex chromosome of Drosophila nasuta nasuta and Drosophila nasuta albomicans." Genome 44, no. 1 (2001): 71–78. http://dx.doi.org/10.1139/g00-100.

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Cellular autoradiography is used to study the transcription patterns of the polytene X chromosomes in Drosophila nasuta nasuta and D. n. albomicans. D. n. nasuta, with 2n = 8, includes a pair of complete heteromorphic sex chromosomes, whereas D. n. albomicans, with 2n = 6, has a pair of metacentric neo-sex chromosomes representing incomplete heteromorphic sex chromosomes. The neo-X chromosome has two euchromatic arms, one representing the ancestral X while the other represents the ancestral autosome 3 chromosomes. The metacentric neo-Y chromosome has one arm with a complete heterochromatic anc
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40

Xu, Wannan, Taiyue Li, Jiahui Li, et al. "The First Identification of Homomorphic XY Sex Chromosomes by Integrating Cytogenetic and Transcriptomic Approaches in Plestiodon elegans (Scincidae)." Genes 15, no. 6 (2024): 664. http://dx.doi.org/10.3390/genes15060664.

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The sex chromosomes of skinks are usually poorly differentiated and hardly distinguished by cytogenetic methods. Therefore, identifying sex chromosomes in species lacking easily recognizable heteromorphic sex chromosomes is necessary to fully understand sex chromosome diversity. In this paper, we employed cytogenetics, sex quantification of genes, and transcriptomic approaches to characterize the sex chromosomes in Plestiodon elegans. Cytogenetic examination of metaphases revealed a diploid number of 2n = 26, consisting of 12 macrochromosomes and 14 microchromosomes, with no significant hetero
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41

Pucci, Marcela B., Patricia Barbosa, Viviane Nogaroto, et al. "Chromosomal Spreading of Microsatellites and (TTAGGG)n Sequences in the Characidium zebra and C. gomesi Genomes (Characiformes: Crenuchidae)." Cytogenetic and Genome Research 149, no. 3 (2016): 182–90. http://dx.doi.org/10.1159/000447959.

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Sex chromosome evolution involves the accumulation of repeat sequences such as multigenic families, noncoding repetitive DNA (satellite, minisatellite, and microsatellite), and mobile elements such as transposons and retrotransposons. Most species of Characidium exhibit heteromorphic ZZ/ZW sex chromosomes; the W is characterized by an intense accumulation of repetitive DNA including dispersed satellite DNA sequences and transposable elements. The aim of this study was to analyze the distribution pattern of 18 different tandem repeats, including (GATA)n and (TTAGGG)n, in the genomes of C. zebra
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42

Röder, G., K. E. Linsenmair, I. Nanda, and M. Schmid. "On sex determination in the Turkish desert woodlouse Hemilepistus elongatus (Crustacea, Isopoda, Oniscidea): searching for sex chromosomes and for sex-specific differences in simple DNA repeats." Genome 39, no. 4 (1996): 818–21. http://dx.doi.org/10.1139/g96-103.

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The karyotype of male and female Hemilepistus elongatus was investigated by means of C-banding. The diploid chromosome number in both sexes is 2n = 50. By scrutinizing general morphology and localization of the constitutive heterochromatin, no heteromorphic sex chromosomes were found. All chromosome pairs in males are well paired during diakinesis. Hybridization of genomic DNA with (GACA)4 and (GATA)4 oligonucleotides revealed no sex-specific patterns. Key words : karyotype, C-banding, sex determination, simple DNA-repeats, Isopoda.
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43

Ngo Ngwe, Florence, and Sonja Siljak-Yakovlev. "Sex Determination in Dioscorea dumetorum: Evidence of Heteromorphic Sex Chromosomes and Sex-Linked NORs." Plants 12, no. 2 (2023): 228. http://dx.doi.org/10.3390/plants12020228.

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Yams (Dioscorea spp.) are a pantropical genus located worldwide that constitute an important source of nutrients and pharmaceutical substances. Some Dioscorea crop species are widely grown in West Africa. One species that is mainly cultivated in Cameroon is Dioscorea dumetorum. This is a dioecious root crop whose sex-determining system was unknown until now. To address the possible presence of sex chromosomes in D. dumetorum, we performed a karyotype characterization of male and female individuals using classical and molecular cytogenetic approaches. It was determined that 2n = 40 was the most
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44

Ciupercescu, D. D., J. Veuskens, A. Mouras, D. Ye, M. Briquet, and I. Negrutiu. "Karyotyping Melandrium album, a dioecious plant with heteromorphic sex chromosomes." Genome 33, no. 4 (1990): 556–62. http://dx.doi.org/10.1139/g90-082.

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Mitotic metaphase chromosomes of Melandrium album obtained from root protoplasts were studied. Morphologically, the chromosomes were either metacentrics or submetacentrics. They were classified into three distinct groups: group A comprising six pairs of autosomal metacentrics, group B comprising five pairs of autosomal submetacentrics, and the sex chromosomes: X and Y. The X chromosome is a metacentric (r = 1.44), which accounts for more than 14% of the genome. The Y chromosome is a metacentric with, virtually, equal arms (r = 1.09) and accounts for 21% of the genome, being the largest of the
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45

Green, David M. "Muller's Ratchet and the evolution of supernumerary chromosomes." Genome 33, no. 6 (1990): 818–24. http://dx.doi.org/10.1139/g90-123.

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Supernumerary chromosomes arise from portions of the normal chromosome complement through nondisjunction, fragmentation, or other mechanisms. Once present in the genome, they are subject to virtually the same genetic conditions that affect the evolutionary degeneration of heteromorphic sex chromosomes. Y or W chromosomes occur only in the presence of X or Z chromosomes, respectively, just as supernumeraries never occur except in the presence of the complete regular karyotype containing their progenitor sequences. Thus, mechanisms that can account for the evolution of sex-chromosome heteromorph
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46

Schmid, Michael, Claus Steinlein, Cassia F. Yano, and Marcelo B. Cioffi. "Hypermethylated Chromosome Regions in Nine Fish Species with Heteromorphic Sex Chromosomes." Cytogenetic and Genome Research 147, no. 2-3 (2015): 169–78. http://dx.doi.org/10.1159/000444067.

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Sites and amounts of 5-methylcytosine (5-MeC)-rich chromosome regions were detected in the karyotypes of 9 Brazilian species of Characiformes fishes by indirect immunofluorescence using a monoclonal anti-5-MeC antibody. These species, belonging to the genera Leporinus, Triportheus and Hoplias, are characterized by highly differentiated and heteromorphic ZW and XY sex chromosomes. In all species, the hypermethylated regions are confined to constitutive heterochromatin. The number and chromosome locations of hypermethylated heterochromatic regions in the karyotypes are constant and species-speci
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47

Joron, M., and A. Whibley. "Stripes, sex and sparrows: what processes underlie heteromorphic chromosome evolution?" Heredity 106, no. 4 (2010): 531–32. http://dx.doi.org/10.1038/hdy.2010.106.

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48

Akagi, Takashi, Naoko Fujita, Kenta Shirasawa, et al. "Rapid and dynamic evolution of a giant Y chromosome in Silene latifolia." Science 387, no. 6734 (2025): 637–43. https://doi.org/10.1126/science.adk9074.

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Some plants have massive sex-linked regions. To test hypotheses about their evolution, we sequenced the genome of Silene latifolia , in which giant heteromorphic sex chromosomes were first discovered in 1923. It has long been known that the Y chromosome consists mainly of a male-specific region that does not recombine with the X chromosome and carries the sex-determining genes and genes with other male functions. However, only with a whole Y chromosome assembly can candidate genes be validated experimentally and their locations determined and related to the suppression of recombination. We des
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49

Liu, Zuyao, Amy L. Herbert, Yingguang Frank Chan, Marek Kučka, David M. Kingsley, and Catherine L. Peichel. "The fourspine stickleback (Apeltes quadracus) has an XY sex chromosome system with polymorphic inversions on both X and Y chromosomes." PLOS Genetics 21, no. 5 (2025): e1011465. https://doi.org/10.1371/journal.pgen.1011465.

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Teleost fish are well-known for possessing a diversity of sex chromosomes and for undergoing frequent turnovers of these sex chromosomes. However, previous studies have mainly focused on variation between species, while comparatively little attention has been given to sex chromosome polymorphisms within species, which may capture early stages of sex chromosome changes. To better understand the evolution of sex chromosomes, we used the fourspine stickleback (Apeltes quadracus) as a model organism. Previous cytogenetic studies suggested that females of this species possessed a ZW heteromorphic s
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50

Grabowska-Joachimiak, Aleksandra, and Andrzej Joachimiak. "C-banded karyotypes of two Silene species with heteromorphic sex chromosomes." Genome 45, no. 2 (2002): 243–52. http://dx.doi.org/10.1139/g01-143.

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Mitotic metaphase chromosomes of Silene latifolia (white campion) and Silene dioica (red campion) were studied and no substantial differences between the conventional karyotypes of these two species were detected. The classification of chromosomes into three distinct groups proposed for S. latifolia by Ciupercescu and colleagues was considered and discussed. Additionally, a new small satellite on the shorter arm of homobrachial chromosome 5 was found. Giemsa C-banded chromosomes of the two analysed species show many fixed and polymorphic heterochromatic bands, mainly distally and centromerical
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