---
_id: '20833'
abstract:
- lang: eng
text: Sex-chromosome systems are highly variable across animals, but how they transition
from one to another is not well understood. Diptera have undergone multiple sex-chromosome
turnovers and expansions while maintaining their general chromosomal content,
which makes them an ideal clade to study such transitions. We analysed more than
100 dipteran whole-genome assemblies and identified 4 new lineages that underwent
sex-chromosome turnover (in addition to the 5 previously reported). We find the
majority of turnovers happened in the group Schizophora, which tend to have fewer
genes on the F element (the chromosome homologous to the ancestral insect X chromosome)
than lower dipterans, a factor previously hypothesized to facilitate turnover.
Most derived X chromosomes have higher GC content than autosomes, consistent with
a high prevalence of male-achiasmy in Diptera. In addition, an excess of gene
movement out of the X is detected for most of these new X chromosomes, and many
of these moved genes have high testis expression in Drosophila, suggesting that
out-of-X gene movement contributes to the long-term demasculinization of X chromosomes.
article_processing_charge: No
author:
- first_name: Lorena Alexandra
full_name: Layana Franco, Lorena Alexandra
id: 02814589-eb8f-11eb-b029-a70074f3f18f
last_name: Layana Franco
orcid: 0000-0002-1253-6297
- first_name: Melissa A
full_name: Toups, Melissa A
id: 4E099E4E-F248-11E8-B48F-1D18A9856A87
last_name: Toups
orcid: 0000-0002-9752-7380
- first_name: Beatriz
full_name: Vicoso, Beatriz
id: 49E1C5C6-F248-11E8-B48F-1D18A9856A87
last_name: Vicoso
orcid: 0000-0002-4579-8306
citation:
ama: Layana Franco LA, Toups MA, Vicoso B. Research Data for “Causes and consequences
of sex-chromosome turnovers in Diptera.” 2026. doi:10.15479/AT-ISTA-20833
apa: Layana Franco, L. A., Toups, M. A., & Vicoso, B. (2026). Research Data
for “Causes and consequences of sex-chromosome turnovers in Diptera.” Institute
of Science and Technology Austria. https://doi.org/10.15479/AT-ISTA-20833
chicago: Layana Franco, Lorena Alexandra, Melissa A Toups, and Beatriz Vicoso. “Research
Data for ‘Causes and Consequences of Sex-Chromosome Turnovers in Diptera.’” Institute
of Science and Technology Austria, 2026. https://doi.org/10.15479/AT-ISTA-20833.
ieee: L. A. Layana Franco, M. A. Toups, and B. Vicoso, “Research Data for ‘Causes
and consequences of sex-chromosome turnovers in Diptera.’” Institute of Science
and Technology Austria, 2026.
ista: Layana Franco LA, Toups MA, Vicoso B. 2026. Research Data for ‘Causes and
consequences of sex-chromosome turnovers in Diptera’, Institute of Science and
Technology Austria, 10.15479/AT-ISTA-20833.
mla: Layana Franco, Lorena Alexandra, et al. Research Data for “Causes and Consequences
of Sex-Chromosome Turnovers in Diptera.” Institute of Science and Technology
Austria, 2026, doi:10.15479/AT-ISTA-20833.
short: L.A. Layana Franco, M.A. Toups, B. Vicoso, (2026).
corr_author: '1'
date_created: 2025-12-17T10:10:57Z
date_published: 2026-01-08T00:00:00Z
date_updated: 2026-06-10T09:21:49Z
day: '8'
department:
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doi: 10.15479/AT-ISTA-20833
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success: 1
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has_accepted_license: '1'
keyword:
- Schizophora
- sex chromosomes
- sex-chromosome turnover
- Diptera
- genomic features
- out-of-X movement.
license: https://creativecommons.org/licenses/by/4.0/
month: '01'
oa: 1
oa_version: Published Version
publisher: Institute of Science and Technology Austria
status: public
title: Research Data for 'Causes and consequences of sex-chromosome turnovers in Diptera'
tmp:
image: /images/cc_by.png
legal_code_url: https://creativecommons.org/licenses/by/4.0/legalcode
name: Creative Commons Attribution 4.0 International Public License (CC-BY 4.0)
short: CC BY (4.0)
type: research_data
user_id: 68b8ca59-c5b3-11ee-8790-cd641c68093d
year: '2026'
...
---
_id: '21116'
abstract:
- lang: eng
text: Sex-chromosome systems are highly variable across animals, but how they transition
from one to another is not well understood. Diptera have undergone multiple sex-chromosome
turnovers and expansions while maintaining their general chromosomal content,
which makes them an ideal clade to study such transitions. We analyzed more than
100 dipteran whole-genome assemblies and identified 4 new lineages that underwent
sex-chromosome turnover (in addition to the 5 previously reported). We find that
the majority of turnovers happened in the group Schizophora, which tend to have
fewer genes on Muller element F (the chromosome homologous to the ancestral insect
X chromosome) than lower dipterans, a factor previously hypothesized to facilitate
turnover. Most derived X chromosomes have higher GC content than autosomes, consistent
with a high prevalence of male achiasmy in Diptera. In addition, an excess of
gene movement out of the X is detected for most of these new X chromosomes, and
many of these moved genes have high testis expression in Drosophila, suggesting
that out-of-X gene movement contributes to the long-term demasculinization of
X chromosomes.
article_processing_charge: No
author:
- first_name: Lorena Alexandra
full_name: Layana Franco, Lorena Alexandra
id: 02814589-eb8f-11eb-b029-a70074f3f18f
last_name: Layana Franco
orcid: 0000-0002-1253-6297
- first_name: Melissa A
full_name: Toups, Melissa A
id: 4E099E4E-F248-11E8-B48F-1D18A9856A87
last_name: Toups
orcid: 0000-0002-9752-7380
- first_name: Beatriz
full_name: Vicoso, Beatriz
id: 49E1C5C6-F248-11E8-B48F-1D18A9856A87
last_name: Vicoso
orcid: 0000-0002-4579-8306
citation:
ama: Layana Franco LA, Toups MA, Vicoso B. Research Data for “Causes and consequences
of sex-chromosome turnovers in Diptera.” 2026. doi:10.15479/AT-ISTA-21116
apa: Layana Franco, L. A., Toups, M. A., & Vicoso, B. (2026). Research Data
for “Causes and consequences of sex-chromosome turnovers in Diptera.” Institute
of Science and Technology Austria. https://doi.org/10.15479/AT-ISTA-21116
chicago: Layana Franco, Lorena Alexandra, Melissa A Toups, and Beatriz Vicoso. “Research
Data for ‘Causes and Consequences of Sex-Chromosome Turnovers in Diptera.’” Institute
of Science and Technology Austria, 2026. https://doi.org/10.15479/AT-ISTA-21116.
ieee: L. A. Layana Franco, M. A. Toups, and B. Vicoso, “Research Data for ‘Causes
and consequences of sex-chromosome turnovers in Diptera.’” Institute of Science
and Technology Austria, 2026.
ista: Layana Franco LA, Toups MA, Vicoso B. 2026. Research Data for ‘Causes and
consequences of sex-chromosome turnovers in Diptera’, Institute of Science and
Technology Austria, 10.15479/AT-ISTA-21116.
mla: Layana Franco, Lorena Alexandra, et al. Research Data for “Causes and Consequences
of Sex-Chromosome Turnovers in Diptera.” Institute of Science and Technology
Austria, 2026, doi:10.15479/AT-ISTA-21116.
short: L.A. Layana Franco, M.A. Toups, B. Vicoso, (2026).
corr_author: '1'
date_created: 2026-01-30T11:04:14Z
date_published: 2026-01-02T00:00:00Z
date_updated: 2026-02-12T12:58:00Z
day: '2'
department:
- _id: BeVi
doi: 10.15479/AT-ISTA-21116
file:
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creator: llayanaf
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date_updated: 2026-01-30T11:00:56Z
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relation: main_file
success: 1
file_date_updated: 2026-01-30T11:00:56Z
fulldoi: https://doi.org/10.15479/AT-ISTA-21116
has_accepted_license: '1'
keyword:
- Schizophora
- sex chromosomes
- sex-chromosome turnover
- Diptera
- genomic features
- out-of-X movement.
month: '01'
oa: 1
oa_version: Published Version
publisher: Institute of Science and Technology Austria
status: public
title: Research Data for "Causes and consequences of sex-chromosome turnovers in Diptera"
tmp:
image: /images/cc_by.png
legal_code_url: https://creativecommons.org/licenses/by/4.0/legalcode
name: Creative Commons Attribution 4.0 International Public License (CC-BY 4.0)
short: CC BY (4.0)
type: research_data
user_id: 68b8ca59-c5b3-11ee-8790-cd641c68093d
year: '2026'
...
---
_id: '20780'
abstract:
- lang: eng
text: Sex-chromosome systems are highly variable across animals, but how they transition
from one to another is not well understood. Diptera have undergone multiple sex-chromosome
turnovers and expansions while maintaining their general chromosomal content,
which makes them an ideal clade to study such transitions. We analysed more than
100 dipteran whole-genome assemblies and identified 4 new lineages that underwent
sex-chromosome turnover (in addition to the 5 previously reported). We find the
majority of turnovers happened in the group Schizophora, which tend to have fewer
genes on the F element (the chromosome homologous to the ancestral insect X chromosome)
than lower dipterans, a factor previously hypothesized to facilitate turnover.
Most derived X chromosomes have higher GC content than autosomes, consistent with
a high prevalence of male-achiasmy in Diptera. In addition, an excess of gene
movement out of the X is detected for most of these new X chromosomes, and many
of these moved genes have high testis expression in Drosophila, suggesting that
out-of-X gene movement contributes to the long-term demasculinization of X chromosomes.
acknowledged_ssus:
- _id: ScienComp
article_processing_charge: No
author:
- first_name: Lorena Alexandra
full_name: Layana Franco, Lorena Alexandra
id: 02814589-eb8f-11eb-b029-a70074f3f18f
last_name: Layana Franco
orcid: 0000-0002-1253-6297
- first_name: Melissa A
full_name: Toups, Melissa A
id: 4E099E4E-F248-11E8-B48F-1D18A9856A87
last_name: Toups
orcid: 0000-0002-9752-7380
- first_name: Beatriz
full_name: Vicoso, Beatriz
id: 49E1C5C6-F248-11E8-B48F-1D18A9856A87
last_name: Vicoso
orcid: 0000-0002-4579-8306
citation:
ama: Layana Franco LA, Toups MA, Vicoso B. Causes and consequences of sex-chromosome
turnovers in Diptera. 2025. doi:10.15479/AT-ISTA-20780
apa: Layana Franco, L. A., Toups, M. A., & Vicoso, B. (2025). Causes and consequences
of sex-chromosome turnovers in Diptera. Institute of Science and Technology Austria.
https://doi.org/10.15479/AT-ISTA-20780
chicago: Layana Franco, Lorena Alexandra, Melissa A Toups, and Beatriz Vicoso. “Causes
and Consequences of Sex-Chromosome Turnovers in Diptera.” Institute of Science
and Technology Austria, 2025. https://doi.org/10.15479/AT-ISTA-20780.
ieee: L. A. Layana Franco, M. A. Toups, and B. Vicoso, “Causes and consequences
of sex-chromosome turnovers in Diptera.” Institute of Science and Technology Austria,
2025.
ista: Layana Franco LA, Toups MA, Vicoso B. 2025. Causes and consequences of sex-chromosome
turnovers in Diptera, Institute of Science and Technology Austria, 10.15479/AT-ISTA-20780.
mla: Layana Franco, Lorena Alexandra, et al. Causes and Consequences of Sex-Chromosome
Turnovers in Diptera. Institute of Science and Technology Austria, 2025, doi:10.15479/AT-ISTA-20780.
short: L.A. Layana Franco, M.A. Toups, B. Vicoso, (2025).
corr_author: '1'
date_created: 2025-12-10T23:40:14Z
date_published: 2025-12-01T00:00:00Z
date_updated: 2026-06-10T08:27:48Z
department:
- _id: BeVi
doi: 10.15479/AT-ISTA-20780
file:
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creator: llayanaf
date_created: 2025-12-11T10:52:17Z
date_updated: 2025-12-11T10:52:17Z
file_id: '20800'
file_name: Supplementary_Datasets.zip
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content_type: application/zip
creator: llayanaf
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success: 1
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fulldoi: https://doi.org/10.15479/AT-ISTA-20780
has_accepted_license: '1'
keyword:
- Schizophora
- sex chromosomes
- sex-chromosome turnover
- Diptera
- genomic features
- out-of-X movement.
month: '12'
oa: 1
oa_version: Published Version
publisher: Institute of Science and Technology Austria
status: public
title: Causes and consequences of sex-chromosome turnovers in Diptera
tmp:
image: /images/cc_by.png
legal_code_url: https://creativecommons.org/licenses/by/4.0/legalcode
name: Creative Commons Attribution 4.0 International Public License (CC-BY 4.0)
short: CC BY (4.0)
type: research_data
user_id: 68b8ca59-c5b3-11ee-8790-cd641c68093d
year: '2025'
...
---
OA_embargo: '6'
OA_place: publisher
_id: '18531'
abstract:
- lang: eng
text: "Sex chromosomes and autosomes exhibit very different evolutionary dynamics.\r\nThe
Y chromosome usually degenerates, leaving many X-linked loci hemizygous in\r\nmales.
Since recessive X-linked mutations are always exposed to selection in males,\r\nselection
is more efficient on the X chromosome than on autosomes on recessive\r\nmutations,
leading to faster adaptation on the X chromosome than other genomic\r\nregions,
if beneficial mutations are on average recessive (known as the Faster-X\r\neffect).
In the presence of the functional, but non-recombining gametolog on the Y (as\r\nis
often the case in young non-recombining regions), recessive mutations are\r\nsheltered
from selection on the X chromosome. We model this scenario and show that\r\nthe
efficiency of selection is reduced on diploid X loci due to sheltering by the
Y\r\nchromosome. Reduced efficiency of selection leads to slower adaptation and\r\nincreased
accumulation of deleterious mutations (Slower-X effect). We extended this\r\nmodel
to explore the effect of sex-specific selection on degeneration of sex\r\nchromosomes,
showing theoretically that male-limited genes degenerate on the X\r\nchromosome
and female-biased genes degenerate on the Y chromosome. This\r\nprediction depends
on the effective population size and the mutation rate, explaining\r\nthe variety
of sex chromosome degeneration patterns observed in nature.\r\nTo test for direct
evidence of a Slower-X (or Slower-Z) effect, we analyzed the\r\nZW sex chromosomes
of the flatworm Schistosoma japonicum, which have a very\r\nyoung non-recombining
region with non-degenerated W. Diploid Z-linked genes have\r\nhigher ratios of
non-synonymous to synonymous polymorphisms than autosomal\r\ngenes, supporting
reduced efficiency of selection on the diploid Z region. These results\r\nprovide
evidence of sheltering by the W chromosome, a mechanism that could\r\ncontribute
to Z (X) chromosome degeneration, and illustrate contrasting evolutionary\r\npatterns
in old and young sex chromosome regions. In addition, genes with sexspecific patterns
of expression show opposite patterns of selection in the young\r\n(diploid) and
old (hemizygous) Z, showing the complex manner in which sex-specific selection
shapes the evolutionary patterns of sex chromosomes. "
acknowledged_ssus:
- _id: ScienComp
- _id: CampIT
alternative_title:
- ISTA Thesis
article_processing_charge: No
author:
- first_name: Andrea
full_name: Mrnjavac, Andrea
id: 353FAC84-AE61-11E9-8BFC-00D3E5697425
last_name: Mrnjavac
citation:
ama: Mrnjavac A. Early stages of sex chromosome evolution. 2024. doi:10.15479/at:ista:18531
apa: Mrnjavac, A. (2024). Early stages of sex chromosome evolution. Institute
of Science and Technology Austria. https://doi.org/10.15479/at:ista:18531
chicago: Mrnjavac, Andrea. “Early Stages of Sex Chromosome Evolution.” Institute
of Science and Technology Austria, 2024. https://doi.org/10.15479/at:ista:18531.
ieee: A. Mrnjavac, “Early stages of sex chromosome evolution,” Institute of Science
and Technology Austria, 2024.
ista: Mrnjavac A. 2024. Early stages of sex chromosome evolution. Institute of Science
and Technology Austria.
mla: Mrnjavac, Andrea. Early Stages of Sex Chromosome Evolution. Institute
of Science and Technology Austria, 2024, doi:10.15479/at:ista:18531.
short: A. Mrnjavac, Early Stages of Sex Chromosome Evolution, Institute of Science
and Technology Austria, 2024.
corr_author: '1'
date_created: 2024-11-11T08:40:45Z
date_published: 2024-11-11T00:00:00Z
date_updated: 2026-07-29T08:36:28Z
day: '11'
ddc:
- '576'
degree_awarded: PhD
department:
- _id: GradSch
- _id: BeVi
doi: 10.15479/at:ista:18531
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title: Early stages of sex chromosome evolution
file_date_updated: 2025-05-11T22:30:04Z
fulldoi: https://doi.org/10.15479/at:ista:18531
has_accepted_license: '1'
keyword:
- Sex chromosomes
- evolution
- selection
- sheltering
language:
- iso: eng
license: https://creativecommons.org/licenses/by-nc-nd/4.0/
month: '11'
oa: 1
oa_version: Published Version
page: '181'
publication_identifier:
issn:
- 2663-337X
publication_status: published
publisher: Institute of Science and Technology Austria
related_material:
record:
- id: '12521'
relation: part_of_dissertation
status: public
- id: '18549'
relation: part_of_dissertation
status: public
status: public
supervisor:
- first_name: Beatriz
full_name: Vicoso, Beatriz
id: 49E1C5C6-F248-11E8-B48F-1D18A9856A87
last_name: Vicoso
orcid: 0000-0002-4579-8306
title: Early stages of sex chromosome evolution
tmp:
image: /images/cc_by_nc_nd.png
legal_code_url: https://creativecommons.org/licenses/by-nc-nd/4.0/legalcode
name: Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International
(CC BY-NC-ND 4.0)
short: CC BY-NC-ND (4.0)
type: dissertation
user_id: ba8df636-2132-11f1-aed0-ed93e2281fdd
year: '2024'
...
---
_id: '10167'
abstract:
- lang: eng
text: Schistosomes, the human parasites responsible for snail fever, are female-heterogametic.
Different parts of their ZW sex chromosomes have stopped recombining in distinct
lineages, creating “evolutionary strata” of various ages. Although the Z-chromosome
is well characterized at the genomic and molecular level, the W-chromosome has
remained largely unstudied from an evolutionary perspective, as only a few W-linked
genes have been detected outside of the model species Schistosoma mansoni. Here,
we characterize the gene content and evolution of the W-chromosomes of S. mansoni
and of the divergent species S. japonicum. We use a combined RNA/DNA k-mer based
pipeline to assemble around 100 candidate W-specific transcripts in each of the
species. About half of them map to known protein coding genes, the majority homologous
to S. mansoni Z-linked genes. We perform an extended analysis of the evolutionary
strata present in the two species (including characterizing a previously undetected
young stratum in S. japonicum) to infer patterns of sequence and expression evolution
of W-linked genes at different time points after recombination was lost. W-linked
genes show evidence of degeneration, including high rates of protein evolution
and reduced expression. Most are found in young lineage-specific strata, with
only a few high expression ancestral W-genes remaining, consistent with the progressive
erosion of nonrecombining regions. Among these, the splicing factor u2af2 stands
out as a promising candidate for primary sex determination, opening new avenues
for understanding the molecular basis of the reproductive biology of this group.
acknowledged_ssus:
- _id: ScienComp
acknowledgement: The authors thank IT support at IST Austria for providing an optimal
environment for bioinformatic analyses. This work was supported by an Austrian Science
Foundation FWF grant (Project P28842) to B.V.
article_processing_charge: No
article_type: original
author:
- first_name: Marwan N
full_name: Elkrewi, Marwan N
id: 0B46FACA-A8E1-11E9-9BD3-79D1E5697425
last_name: Elkrewi
orcid: 0000-0002-5328-7231
- first_name: Mikhail A.
full_name: Moldovan, Mikhail A.
id: c8bb7f32-3315-11ec-b58b-e5950e6c14a0
last_name: Moldovan
orcid: 0000-0002-8876-6494
- first_name: Marion A L
full_name: Picard, Marion A L
id: 2C921A7A-F248-11E8-B48F-1D18A9856A87
last_name: Picard
orcid: 0000-0002-8101-2518
- first_name: Beatriz
full_name: Vicoso, Beatriz
id: 49E1C5C6-F248-11E8-B48F-1D18A9856A87
last_name: Vicoso
orcid: 0000-0002-4579-8306
citation:
ama: Elkrewi MN, Moldovan MA, Picard MAL, Vicoso B. Schistosome W-linked genes inform
temporal dynamics of sex chromosome evolution and suggest candidate for sex determination.
Molecular Biology and Evolution. 2021;138(12):5345-5358. doi:10.1093/molbev/msab178
apa: Elkrewi, M. N., Moldovan, M. A., Picard, M. A. L., & Vicoso, B. (2021).
Schistosome W-linked genes inform temporal dynamics of sex chromosome evolution
and suggest candidate for sex determination. Molecular Biology and Evolution.
Oxford University Press. https://doi.org/10.1093/molbev/msab178
chicago: Elkrewi, Marwan N, Mikhail A. Moldovan, Marion A L Picard, and Beatriz
Vicoso. “Schistosome W-Linked Genes Inform Temporal Dynamics of Sex Chromosome
Evolution and Suggest Candidate for Sex Determination.” Molecular Biology and
Evolution. Oxford University Press, 2021. https://doi.org/10.1093/molbev/msab178.
ieee: M. N. Elkrewi, M. A. Moldovan, M. A. L. Picard, and B. Vicoso, “Schistosome
W-linked genes inform temporal dynamics of sex chromosome evolution and suggest
candidate for sex determination,” Molecular Biology and Evolution, vol.
138, no. 12. Oxford University Press, pp. 5345–58, 2021.
ista: Elkrewi MN, Moldovan MA, Picard MAL, Vicoso B. 2021. Schistosome W-linked
genes inform temporal dynamics of sex chromosome evolution and suggest candidate
for sex determination. Molecular Biology and Evolution. 138(12), 5345–58.
mla: Elkrewi, Marwan N., et al. “Schistosome W-Linked Genes Inform Temporal Dynamics
of Sex Chromosome Evolution and Suggest Candidate for Sex Determination.” Molecular
Biology and Evolution, vol. 138, no. 12, Oxford University Press, 2021, pp.
5345–58, doi:10.1093/molbev/msab178.
short: M.N. Elkrewi, M.A. Moldovan, M.A.L. Picard, B. Vicoso, Molecular Biology
and Evolution 138 (2021) 5345–58.
corr_author: '1'
das_tickbox: '1'
date_created: 2021-10-21T07:49:12Z
date_published: 2021-06-19T00:00:00Z
date_updated: 2026-09-13T22:30:29Z
day: '19'
ddc:
- '610'
department:
- _id: BeVi
doi: 10.1093/molbev/msab178
external_id:
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pmid:
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intvolume: ' 138'
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issue: '12'
keyword:
- sex chromosomes
- evolutionary strata
- W-linked gene
- sex determining gene
- schistosome parasites
language:
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month: '06'
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call_identifier: FWF
grant_number: P28842-B22
name: Sex chromosome evolution under male- and female- heterogamety
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title: Schistosome W-linked genes inform temporal dynamics of sex chromosome evolution
and suggest candidate for sex determination
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