---
OA_place: publisher
_id: '20694'
abstract:
- lang: eng
  text: "Understanding the mechanisms underlying speciation is a central aim of evolutionary
    biology.\r\nA persistent challenge in the field is to identify loci that contribute
    to reproductive isolation,\r\nwhile disentangling signals of selection from demography,
    linkage and intrinsic genomic\r\nfeatures. Traditional population genomic approaches
    that rely on site-based statistics in\r\narbitrary fixed windows face inherent
    limitations, as they conflate historical and\r\ncontemporary processes of divergence
    and overlook haplotype structure. Recent advances in\r\nwhole-genome sequencing
    and methods to infer ancestral recombination graphs (ARGs) now\r\noffer the opportunity
    to study genealogical relationships explicitly, revealing how lineages\r\ncoalesce
    and recombine through time. By directly analysing haplotype clustering by species\r\nor
    phenotype and their patterns of coalescence, ARG-based methods show promise for\r\ndiagnosing
    sweeps, identifying barrier loci maintained under divergent selection amid gene\r\nflow,
    and tracing their evolutionary history.\r\nIn this thesis, I explore the utility
    of genealogical approaches for studying species\r\ndivergence. In chapter 2, I
    propose a conceptual framework for defining haplotype blocks\r\nthrough the structure
    of the ARG, using simulations and empirical data to highlight how\r\ngenealogical
    processes generate rich and often overlooked haplotypic patterns.\r\nIn chapter
    3, I examine the genomic basis of a key evolutionary innovation in marine\r\nsnails
    Littorina. These snails offer a unique opportunity to study an innovation because
    they\r\ninclude a very recent transition from egg-laying to live bearing, yet
    snails with the different\r\nreproductive modes are not reciprocally monophyletic.
    I exploited this by using topology\r\nclustering in ARG-derived local genealogical
    trees to pinpoint narrow genomic regions or\r\nhaplotype blocks that carry swept
    alleles, thus revealing that the transition from egg-laying\r\nto live-bearing
    involves multiple, live-bearer-specific sweeps.\r\nChapter 4 establishes a population-scale,
    phased genomic resource for Antirrhinum\r\nmajus, using cost-effective haplotagging,
    then optimizes imputation from low-coverage data\r\nagainst high-accuracy KASP
    sequencing to maximize sequence completeness with modest\r\naccuracy trade-offs
    against a traditional short-read sequence pipeline. A hybrid phasing\r\nstrategy
    combines molecular phasing with statistical phasing to generate phased whole\r\ngenome
    sequences of 1084 Antirrhinum individuals at a fraction of long-read sequencing\r\ncosts.\r\nIn
    chapter 5, I analyse hybridising populations from two replicate hybrid zones to
    find\r\na parallel genetic basis of flower colour, amidst the noise in genomic
    differentiation landscape\r\ndriven by variation in demographic history. While
    outlier genome scans of FST failed to dissect\r\nthe causes of differentiation,
    ARG-based topology clustering revealed a reuse of colour\r\nassociated haplotypes
    across hybrid zones. In addition to the biological insight, this chapter\r\nalso
    presents a comparison of the latest ARG inference tools, showing that signals
    of\r\nAbstract\r\nviii\r\ntopological clustering qualitatively agree between methods,
    despite differences in the tree\r\nsequences.\r\nNext, in chapter 6, by leveraging
    ~1000 individuals in one of the hybrid zones, I\r\nintegrated genome-wide association
    studies of floral pigmentation with genealogical\r\ninference, to test for additional
    colour loci, and confirm the effect of previously described loci.\r\nThis work
    demonstrates that flower colour variation is driven by a small number of large
    effect\r\nloci, while also hinting at the presence of a new candidate regulatory
    factor.\r\nFinally in chapter 7, in a preliminary analysis, I begin to dissect
    the genomic island of\r\nspeciation around Rosea/Eluta to understand its evolutionary
    origins. My results show that it\r\nconsists of 5 highly divergent loci, each
    of which is associated with flower colour. Using\r\npatterns of coalescence in
    genealogical trees, I find evidence of staggered selective sweeps\r\nand a persistent
    localized barrier to gene flow within an otherwise permeable genome.\r\nTogether,
    these chapters add to the increasing pool of studies using genealogical\r\napproaches
    to complement and extend site-based statistics to use haplotype structures in\r\nspeciation
    research. By tracking haplotypes directly and connecting genealogical clustering
    to\r\npopulation processes, ARG-based inference promises to provide new insights
    into how local\r\nselective pressures, demographic history, and long-term barriers
    interact to shape the\r\ngenomic architecture of divergence. By underscoring the
    value of ARGs in revealing the finescale origins and maintenance of biodiversity,
    this thesis presents cautious optimism about\r\nthe benefits of using genealogical
    inference to learn more than what site-based statistics\r\ncould tell us."
acknowledged_ssus:
- _id: ScienComp
alternative_title:
- ISTA Thesis
article_processing_charge: No
author:
- first_name: Arka
  full_name: Pal, Arka
  id: 6AAB2240-CA9A-11E9-9C1A-D9D1E5697425
  last_name: Pal
  orcid: 0000-0002-4530-8469
citation:
  ama: Pal A. Using genealogies to study the genomic basis of species divergence.
    2025. doi:<a href="https://doi.org/10.15479/AT-ISTA-20694">10.15479/AT-ISTA-20694</a>
  apa: Pal, A. (2025). <i>Using genealogies to study the genomic basis of species
    divergence</i>. Institute of Science and Technology Austria. <a href="https://doi.org/10.15479/AT-ISTA-20694">https://doi.org/10.15479/AT-ISTA-20694</a>
  chicago: Pal, Arka. “Using Genealogies to Study the Genomic Basis of Species Divergence.”
    Institute of Science and Technology Austria, 2025. <a href="https://doi.org/10.15479/AT-ISTA-20694">https://doi.org/10.15479/AT-ISTA-20694</a>.
  ieee: A. Pal, “Using genealogies to study the genomic basis of species divergence,”
    Institute of Science and Technology Austria, 2025.
  ista: Pal A. 2025. Using genealogies to study the genomic basis of species divergence.
    Institute of Science and Technology Austria.
  mla: Pal, Arka. <i>Using Genealogies to Study the Genomic Basis of Species Divergence</i>.
    Institute of Science and Technology Austria, 2025, doi:<a href="https://doi.org/10.15479/AT-ISTA-20694">10.15479/AT-ISTA-20694</a>.
  short: A. Pal, Using Genealogies to Study the Genomic Basis of Species Divergence,
    Institute of Science and Technology Austria, 2025.
corr_author: '1'
date_created: 2025-11-25T13:19:11Z
date_published: 2025-11-25T00:00:00Z
date_updated: 2026-04-28T13:20:36Z
day: '25'
ddc:
- '576'
- '578'
degree_awarded: PhD
department:
- _id: GradSch
- _id: NiBa
doi: 10.15479/AT-ISTA-20694
file:
- access_level: open_access
  checksum: 7a10a738d58524aebb5dcbd9b34c21c5
  content_type: application/pdf
  creator: apal
  date_created: 2025-12-01T13:53:36Z
  date_updated: 2026-03-01T23:30:03Z
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  creator: apal
  date_created: 2025-12-01T13:53:39Z
  date_updated: 2026-03-01T23:30:03Z
  embargo_to: open_access
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  file_name: 2025_Pal_Arka_Thesis.docx
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file_date_updated: 2026-03-01T23:30:03Z
has_accepted_license: '1'
language:
- iso: eng
license: https://creativecommons.org/licenses/by-nc-nd/4.0/
month: '11'
oa: 1
oa_version: Published Version
page: '268'
project:
- _id: bd6958e0-d553-11ed-ba76-86eba6a76c00
  grant_number: '101055327'
  name: Understanding the evolution of continuous genomes
- _id: 05959E1C-7A3F-11EA-A408-12923DDC885E
  grant_number: P32166
  name: Snapdragon Speciation
publication_identifier:
  issn:
  - 2663-337X
publication_status: published
publisher: Institute of Science and Technology Austria
related_material:
  record:
  - id: '12159'
    relation: part_of_dissertation
    status: public
  - id: '14796'
    relation: part_of_dissertation
    status: public
  - id: '20190'
    relation: part_of_dissertation
    status: public
status: public
supervisor:
- first_name: Nicholas H
  full_name: Barton, Nicholas H
  id: 4880FE40-F248-11E8-B48F-1D18A9856A87
  last_name: Barton
  orcid: 0000-0002-8548-5240
title: Using genealogies to study the genomic basis of species divergence
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: '2025'
...
---
OA_place: publisher
OA_type: hybrid
PlanS_conform: '1'
_id: '20190'
abstract:
- lang: eng
  text: 'A major goal of speciation research is identifying loci that underpin barriers
    to gene flow. Population genomics takes a ‘bottom-up’ approach, scanning the genome
    for molecular signatures of processes that drive or maintain divergence. However,
    interpreting the ‘genomic landscape’ of speciation is complicated, because genome
    scans conflate multiple processes, most of which are not informative about gene
    flow. However, studying replicated population contrasts, including multiple incidences
    of secondary contact, can strengthen inferences. In this paper, we use linked-read
    sequencing (haplotagging), FST scans and genealogical methods to characterise
    the genomic landscape associated with replicate hybrid zone formation. We studied
    two flower colour varieties of the common snapdragon, Antirrhinum majus subspecies
    majus, that form secondary hybrid zones in multiple independent valleys in the
    Pyrenees. Consistent with past work, we found very low differentiation at one
    well-studied zone (Planoles). However, at a second zone (Avellanet), we found
    stronger differentiation and greater heterogeneity, which we argue is due to differences
    in the amount of introgression following secondary contact. Topology weighting
    of genealogical trees identified loci where haplotype diversity was associated
    with the two snapdragon varieties. Two of the strongest associations were at previously
    identified flower colour loci: Flavia, that affects yellow pigmentation, and Rosea/Eluta,
    two linked loci that affect magenta pigmentation. Preliminary analysis of coalescence
    times provides additional evidence for selective sweeps at these loci and barriers
    to gene flow. Our study highlights the impact of demographic history on the differentiation
    landscape, emphasising the need to distinguish between historical divergence and
    recent introgression.'
acknowledged_ssus:
- _id: ScienComp
acknowledgement: 'We thank ESEB Godfrey Hewitt Mobility Award for supporting AP’s
  research stay at UC Davis. We thank Tom Ellis, Parvathy Surendranadh, and other
  Barton Group and Coop Lab members for stimulating discussions. We are grateful to
  all the interns and volunteers who have helped us with fieldwork. We thank Eva Salmerón
  Mateu for her assistance in fieldwork logistics at the field station, El Serrat.
  We are grateful to Enrico Coen and his research group for providing the Antirrhinum
  molle PoolSeq data used in the allele polarisation. We are also thankful to Enrico
  Coen and Cristophe Thébaud for discovering the Avellanet hybrid zone, followed up
  with sampling led by D.L.F. in 2017. The study was supported by Austrian Science
  Fund (FWF) Grant (Snapdragon Speciation P32166, awarded to D.L.F.); ERC (Advanced
  Grant HaplotypeStructure 101055327, awarded to NHB); ERC (POC Grant 101069216, awarded
  to Y.F.C.) and the National Institutes of Health (NIH R35 GM136290, awarded to G.C.).
  Y.F.C. was supported by the Max Planck Society. Computing infrastructure for bioinformatics
  and analyses was provided by ISTA High Performance Cluster. '
article_number: e70067
article_processing_charge: Yes (via OA deal)
article_type: original
author:
- first_name: Arka
  full_name: Pal, Arka
  id: 6AAB2240-CA9A-11E9-9C1A-D9D1E5697425
  last_name: Pal
  orcid: 0000-0002-4530-8469
- first_name: Daria
  full_name: Shipilina, Daria
  id: 428A94B0-F248-11E8-B48F-1D18A9856A87
  last_name: Shipilina
  orcid: 0000-0002-1145-9226
- first_name: Alan
  full_name: Le Moan, Alan
  last_name: Le Moan
- first_name: Adrian J.
  full_name: Mcnairn, Adrian J.
  last_name: Mcnairn
- first_name: Jennifer K.
  full_name: Grenier, Jennifer K.
  last_name: Grenier
- first_name: Marek
  full_name: Kucka, Marek
  last_name: Kucka
- first_name: Graham
  full_name: Coop, Graham
  last_name: Coop
- first_name: Yingguang Frank
  full_name: Chan, Yingguang Frank
  last_name: Chan
- first_name: Nicholas H
  full_name: Barton, Nicholas H
  id: 4880FE40-F248-11E8-B48F-1D18A9856A87
  last_name: Barton
  orcid: 0000-0002-8548-5240
- first_name: David
  full_name: Field, David
  id: 419049E2-F248-11E8-B48F-1D18A9856A87
  last_name: Field
  orcid: 0000-0002-4014-8478
- first_name: Sean
  full_name: Stankowski, Sean
  id: 43161670-5719-11EA-8025-FABC3DDC885E
  last_name: Stankowski
citation:
  ama: Pal A, Shipilina D, Le Moan A, et al. Genealogical analysis of replicate flower
    colour hybrid zones in Antirrhinum. <i>Molecular Ecology</i>. 2025;34(22). doi:<a
    href="https://doi.org/10.1111/mec.70067">10.1111/mec.70067</a>
  apa: Pal, A., Shipilina, D., Le Moan, A., Mcnairn, A. J., Grenier, J. K., Kucka,
    M., … Stankowski, S. (2025). Genealogical analysis of replicate flower colour
    hybrid zones in Antirrhinum. <i>Molecular Ecology</i>. Wiley. <a href="https://doi.org/10.1111/mec.70067">https://doi.org/10.1111/mec.70067</a>
  chicago: Pal, Arka, Daria Shipilina, Alan Le Moan, Adrian J. Mcnairn, Jennifer K.
    Grenier, Marek Kucka, Graham Coop, et al. “Genealogical Analysis of Replicate
    Flower Colour Hybrid Zones in Antirrhinum.” <i>Molecular Ecology</i>. Wiley, 2025.
    <a href="https://doi.org/10.1111/mec.70067">https://doi.org/10.1111/mec.70067</a>.
  ieee: A. Pal <i>et al.</i>, “Genealogical analysis of replicate flower colour hybrid
    zones in Antirrhinum,” <i>Molecular Ecology</i>, vol. 34, no. 22. Wiley, 2025.
  ista: Pal A, Shipilina D, Le Moan A, Mcnairn AJ, Grenier JK, Kucka M, Coop G, Chan
    YF, Barton NH, Field D, Stankowski S. 2025. Genealogical analysis of replicate
    flower colour hybrid zones in Antirrhinum. Molecular Ecology. 34(22), e70067.
  mla: Pal, Arka, et al. “Genealogical Analysis of Replicate Flower Colour Hybrid
    Zones in Antirrhinum.” <i>Molecular Ecology</i>, vol. 34, no. 22, e70067, Wiley,
    2025, doi:<a href="https://doi.org/10.1111/mec.70067">10.1111/mec.70067</a>.
  short: A. Pal, D. Shipilina, A. Le Moan, A.J. Mcnairn, J.K. Grenier, M. Kucka, G.
    Coop, Y.F. Chan, N.H. Barton, D. Field, S. Stankowski, Molecular Ecology 34 (2025).
corr_author: '1'
date_created: 2025-08-17T22:01:37Z
date_published: 2025-11-01T00:00:00Z
date_updated: 2026-05-05T22:31:12Z
day: '01'
ddc:
- '570'
department:
- _id: NiBa
doi: 10.1111/mec.70067
external_id:
  isi:
  - '001546622100001'
file:
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  date_created: 2026-01-05T13:47:47Z
  date_updated: 2026-01-05T13:47:47Z
  file_id: '20958'
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  success: 1
file_date_updated: 2026-01-05T13:47:47Z
has_accepted_license: '1'
intvolume: '        34'
isi: 1
issue: '22'
language:
- iso: eng
month: '11'
oa: 1
oa_version: Published Version
project:
- _id: 05959E1C-7A3F-11EA-A408-12923DDC885E
  grant_number: P32166
  name: Snapdragon Speciation
- _id: bd6958e0-d553-11ed-ba76-86eba6a76c00
  grant_number: '101055327'
  name: Understanding the evolution of continuous genomes
publication: Molecular Ecology
publication_identifier:
  eissn:
  - 1365-294X
  issn:
  - 0962-1083
publication_status: published
publisher: Wiley
quality_controlled: '1'
related_material:
  link:
  - description: News on ISTA website
    relation: press_release
    url: https://ista.ac.at/en/news/snapdragon-secrets/
  record:
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    relation: dissertation_contains
    status: public
scopus_import: '1'
status: public
title: Genealogical analysis of replicate flower colour hybrid zones in Antirrhinum
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: journal_article
user_id: ba8df636-2132-11f1-aed0-ed93e2281fdd
volume: 34
year: '2025'
...
---
_id: '14850'
abstract:
- lang: eng
  text: Elaborate sexual signals are thought to have evolved and be maintained to
    serve as honest indicators of signaller quality. One measure of quality is health,
    which can be affected by parasite infection. Cnemaspis mysoriensis is a diurnal
    gecko that is often infested with ectoparasites in the wild, and males of this
    species express visual (coloured gular patches) and chemical (femoral gland secretions)
    traits that receivers could assess during social interactions. In this paper,
    we tested whether ectoparasites affect individual health, and whether signal quality
    is an indicator of ectoparasite levels. In wild lizards, we found that ectoparasite
    level was negatively correlated with body condition in both sexes. Moreover, some
    characteristics of both visual and chemical traits in males were strongly associated
    with ectoparasite levels. Specifically, males with higher ectoparasite levels
    had yellow gular patches with lower brightness and chroma, and chemical secretions
    with a lower proportion of aromatic compounds. We then determined whether ectoparasite
    levels in males influence female behaviour. Using sequential choice trials, wherein
    females were provided with either the visual or the chemical signals of wild-caught
    males that varied in ectoparasite level, we found that only chemical secretions
    evoked an elevated female response towards less parasitised males. Simultaneous
    choice trials in which females were exposed to the chemical secretions from males
    that varied in parasite level further confirmed a preference for males with lower
    parasites loads. Overall, we find that although health (body condition) or ectoparasite
    load can be honestly advertised through multiple modalities, the parasite-mediated
    female response is exclusively driven by chemical signals.</jats:p>
acknowledgement: "We thank Anuradha Batabyal and Shakilur Kabir for scientific discussions,
  and help with sampling and colour analyses. We thank Muralidhar and the central
  LCMS facility of the IISc for their technical support with the GCMS.\r\nResearch
  funding was provided by the Department of Science and Technology Fund for Improvement
  of S&T Infrastructure (DST-FIST), the Department of Biotechnology-Indian Institute
  of Science (DBT-IISc) partnership program and a Science and Engineering Research
  Board (SERB) grant to M.T. (EMR/2017/002228). Open Access funding provided by Indian
  Institute of Science. Deposited in PMC for immediate release."
article_number: jeb246217
article_processing_charge: Yes (via OA deal)
article_type: original
author:
- first_name: Arka
  full_name: Pal, Arka
  id: 6AAB2240-CA9A-11E9-9C1A-D9D1E5697425
  last_name: Pal
  orcid: 0000-0002-4530-8469
- first_name: Mihir
  full_name: Joshi, Mihir
  last_name: Joshi
- first_name: Maria
  full_name: Thaker, Maria
  last_name: Thaker
citation:
  ama: Pal A, Joshi M, Thaker M. Too much information? Males convey parasite levels
    using more signal modalities than females utilise. <i>Journal of Experimental
    Biology</i>. 2024;227(1). doi:<a href="https://doi.org/10.1242/jeb.246217">10.1242/jeb.246217</a>
  apa: Pal, A., Joshi, M., &#38; Thaker, M. (2024). Too much information? Males convey
    parasite levels using more signal modalities than females utilise. <i>Journal
    of Experimental Biology</i>. The Company of Biologists. <a href="https://doi.org/10.1242/jeb.246217">https://doi.org/10.1242/jeb.246217</a>
  chicago: Pal, Arka, Mihir Joshi, and Maria Thaker. “Too Much Information? Males
    Convey Parasite Levels Using More Signal Modalities than Females Utilise.” <i>Journal
    of Experimental Biology</i>. The Company of Biologists, 2024. <a href="https://doi.org/10.1242/jeb.246217">https://doi.org/10.1242/jeb.246217</a>.
  ieee: A. Pal, M. Joshi, and M. Thaker, “Too much information? Males convey parasite
    levels using more signal modalities than females utilise,” <i>Journal of Experimental
    Biology</i>, vol. 227, no. 1. The Company of Biologists, 2024.
  ista: Pal A, Joshi M, Thaker M. 2024. Too much information? Males convey parasite
    levels using more signal modalities than females utilise. Journal of Experimental
    Biology. 227(1), jeb246217.
  mla: Pal, Arka, et al. “Too Much Information? Males Convey Parasite Levels Using
    More Signal Modalities than Females Utilise.” <i>Journal of Experimental Biology</i>,
    vol. 227, no. 1, jeb246217, The Company of Biologists, 2024, doi:<a href="https://doi.org/10.1242/jeb.246217">10.1242/jeb.246217</a>.
  short: A. Pal, M. Joshi, M. Thaker, Journal of Experimental Biology 227 (2024).
corr_author: '1'
date_created: 2024-01-22T08:14:49Z
date_published: 2024-01-10T00:00:00Z
date_updated: 2025-09-04T11:50:21Z
day: '10'
ddc:
- '570'
department:
- _id: NiBa
doi: 10.1242/jeb.246217
external_id:
  isi:
  - '001214515700016'
  pmid:
  - '38054353'
file:
- access_level: open_access
  checksum: 136325372f6f45abaa62a71e2d23bfb6
  content_type: application/pdf
  creator: dernst
  date_created: 2024-01-23T12:08:24Z
  date_updated: 2024-01-23T12:08:24Z
  file_id: '14877'
  file_name: 2024_JourExperimBiology_Pal.pdf
  file_size: 594128
  relation: main_file
  success: 1
file_date_updated: 2024-01-23T12:08:24Z
has_accepted_license: '1'
intvolume: '       227'
isi: 1
issue: '1'
keyword:
- Insect Science
- Molecular Biology
- Animal Science and Zoology
- Aquatic Science
- Physiology
- Ecology
- Evolution
- Behavior and Systematics
language:
- iso: eng
month: '01'
oa: 1
oa_version: Published Version
pmid: 1
publication: Journal of Experimental Biology
publication_identifier:
  eissn:
  - 0022-0949
  issn:
  - 1477-9145
publication_status: published
publisher: The Company of Biologists
quality_controlled: '1'
related_material:
  link:
  - relation: software
    url: https://github.com/arka-pal/Cnemaspis-SexualSignaling
scopus_import: '1'
status: public
title: Too much information? Males convey parasite levels using more signal modalities
  than females utilise
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: journal_article
user_id: 317138e5-6ab7-11ef-aa6d-ffef3953e345
volume: 227
year: '2024'
...
---
OA_place: repository
OA_type: green
_id: '14796'
abstract:
- lang: eng
  text: Key innovations are fundamental to biological diversification, but their genetic
    basis is poorly understood. A recent transition from egg-laying to live-bearing
    in marine snails (Littorina spp.) provides the opportunity to study the genetic
    architecture of an innovation that has evolved repeatedly across animals. Individuals
    do not cluster by reproductive mode in a genome-wide phylogeny, but local genealogical
    analysis revealed numerous small genomic regions where all live-bearers carry
    the same core haplotype. Candidate regions show evidence for live-bearer–specific
    positive selection and are enriched for genes that are differentially expressed
    between egg-laying and live-bearing reproductive systems. Ages of selective sweeps
    suggest that live-bearer–specific alleles accumulated over more than 200,000 generations.
    Our results suggest that new functions evolve through the recruitment of many
    alleles rather than in a single evolutionary step.
acknowledgement: "We thank J. Galindo, M. Montaño-Rendón, N. Mikhailova, A. Blakeslee,
  E. Arnason, and P. Kemppainen for providing samples; R. Turney, G. Sotelo, J. Larsson,
  T. Broquet, and S. Loisel for help collecting samples; Science Animated for providing
  the snail cartoons shown in Fig. 1; M. Dunning for help in developing bioinformatic
  pipelines; R. Faria, H. Morales, and V. Sousa for advice; and M. Hahn, J. Slate,
  M. Ravinet, J. Raeymaekers, A. Comeault, and N. Barton for feedback on a draft manuscript.\r\nThis
  work was supported by the Natural Environment Research Council (grant NE/P001610/1
  to R.K.B.), the European Research Council (grant ERC-2015-AdG693030-BARRIERS to
  R.K.B.), the Norwegian Research Council (RCN Project 315287 to A.M.W.), and the
  Swedish Research Council (grant 2020-05385 to E.L.)."
article_processing_charge: No
article_type: original
author:
- first_name: Sean
  full_name: Stankowski, Sean
  id: 43161670-5719-11EA-8025-FABC3DDC885E
  last_name: Stankowski
- first_name: Zuzanna B.
  full_name: Zagrodzka, Zuzanna B.
  last_name: Zagrodzka
- first_name: Martin D.
  full_name: Garlovsky, Martin D.
  last_name: Garlovsky
- first_name: Arka
  full_name: Pal, Arka
  id: 6AAB2240-CA9A-11E9-9C1A-D9D1E5697425
  last_name: Pal
  orcid: 0000-0002-4530-8469
- first_name: Daria
  full_name: Shipilina, Daria
  id: 428A94B0-F248-11E8-B48F-1D18A9856A87
  last_name: Shipilina
  orcid: 0000-0002-1145-9226
- first_name: Diego Fernando
  full_name: Garcia Castillo, Diego Fernando
  id: ae681a14-dc74-11ea-a0a7-c6ef18161701
  last_name: Garcia Castillo
- first_name: Hila
  full_name: Lifchitz, Hila
  id: d6ab5470-2fb3-11ed-8633-986a9b84edac
  last_name: Lifchitz
- first_name: Alan
  full_name: Le Moan, Alan
  last_name: Le Moan
- first_name: Erica
  full_name: Leder, Erica
  last_name: Leder
- first_name: James
  full_name: Reeve, James
  last_name: Reeve
- first_name: Kerstin
  full_name: Johannesson, Kerstin
  last_name: Johannesson
- first_name: Anja M
  full_name: Westram, Anja M
  id: 3C147470-F248-11E8-B48F-1D18A9856A87
  last_name: Westram
  orcid: 0000-0003-1050-4969
- first_name: Roger K.
  full_name: Butlin, Roger K.
  last_name: Butlin
citation:
  ama: Stankowski S, Zagrodzka ZB, Garlovsky MD, et al. The genetic basis of a recent
    transition to live-bearing in marine snails. <i>Science</i>. 2024;383(6678):114-119.
    doi:<a href="https://doi.org/10.1126/science.adi2982">10.1126/science.adi2982</a>
  apa: Stankowski, S., Zagrodzka, Z. B., Garlovsky, M. D., Pal, A., Shipilina, D.,
    Garcia Castillo, D. F., … Butlin, R. K. (2024). The genetic basis of a recent
    transition to live-bearing in marine snails. <i>Science</i>. American Association
    for the Advancement of Science. <a href="https://doi.org/10.1126/science.adi2982">https://doi.org/10.1126/science.adi2982</a>
  chicago: Stankowski, Sean, Zuzanna B. Zagrodzka, Martin D. Garlovsky, Arka Pal,
    Daria Shipilina, Diego Fernando Garcia Castillo, Hila Lifchitz, et al. “The Genetic
    Basis of a Recent Transition to Live-Bearing in Marine Snails.” <i>Science</i>.
    American Association for the Advancement of Science, 2024. <a href="https://doi.org/10.1126/science.adi2982">https://doi.org/10.1126/science.adi2982</a>.
  ieee: S. Stankowski <i>et al.</i>, “The genetic basis of a recent transition to
    live-bearing in marine snails,” <i>Science</i>, vol. 383, no. 6678. American Association
    for the Advancement of Science, pp. 114–119, 2024.
  ista: Stankowski S, Zagrodzka ZB, Garlovsky MD, Pal A, Shipilina D, Garcia Castillo
    DF, Lifchitz H, Le Moan A, Leder E, Reeve J, Johannesson K, Westram AM, Butlin
    RK. 2024. The genetic basis of a recent transition to live-bearing in marine snails.
    Science. 383(6678), 114–119.
  mla: Stankowski, Sean, et al. “The Genetic Basis of a Recent Transition to Live-Bearing
    in Marine Snails.” <i>Science</i>, vol. 383, no. 6678, American Association for
    the Advancement of Science, 2024, pp. 114–19, doi:<a href="https://doi.org/10.1126/science.adi2982">10.1126/science.adi2982</a>.
  short: S. Stankowski, Z.B. Zagrodzka, M.D. Garlovsky, A. Pal, D. Shipilina, D.F.
    Garcia Castillo, H. Lifchitz, A. Le Moan, E. Leder, J. Reeve, K. Johannesson,
    A.M. Westram, R.K. Butlin, Science 383 (2024) 114–119.
corr_author: '1'
date_created: 2024-01-14T23:00:56Z
date_published: 2024-01-05T00:00:00Z
date_updated: 2026-05-05T22:31:12Z
day: '05'
department:
- _id: NiBa
- _id: GradSch
doi: 10.1126/science.adi2982
external_id:
  isi:
  - '001138156400003'
  pmid:
  - '38175895'
intvolume: '       383'
isi: 1
issue: '6678'
language:
- iso: eng
main_file_link:
- open_access: '1'
  url: https://figshare.com/articles/journal_contribution/The_genetic_basis_of_a_recent_transition_to_live-bearing_in_marine_snails/26356054?file=47868241
month: '01'
oa: 1
oa_version: Submitted Version
page: 114-119
pmid: 1
publication: Science
publication_identifier:
  eissn:
  - 1095-9203
publication_status: published
publisher: American Association for the Advancement of Science
quality_controlled: '1'
related_material:
  link:
  - description: News on ISTA Website
    relation: press_release
    url: https://ista.ac.at/en/news/the-snail-or-the-egg/
  record:
  - id: '14812'
    relation: research_data
    status: public
  - id: '20694'
    relation: dissertation_contains
    status: public
scopus_import: '1'
status: public
title: The genetic basis of a recent transition to live-bearing in marine snails
type: journal_article
user_id: 317138e5-6ab7-11ef-aa6d-ffef3953e345
volume: 383
year: '2024'
...
---
_id: '12159'
abstract:
- lang: eng
  text: The term “haplotype block” is commonly used in the developing field of haplotype-based
    inference methods. We argue that the term should be defined based on the structure
    of the Ancestral Recombination Graph (ARG), which contains complete information
    on the ancestry of a sample. We use simulated examples to demonstrate key features
    of the relationship between haplotype blocks and ancestral structure, emphasizing
    the stochasticity of the processes that generate them. Even the simplest cases
    of neutrality or of a “hard” selective sweep produce a rich structure, often missed
    by commonly used statistics. We highlight a number of novel methods for inferring
    haplotype structure, based on the full ARG, or on a sequence of trees, and illustrate
    how they can be used to define haplotype blocks using an empirical data set. While
    the advent of new, computationally efficient methods makes it possible to apply
    these concepts broadly, they (and additional new methods) could benefit from adding
    features to explore haplotype blocks, as we define them. Understanding and applying
    the concept of the haplotype block will be essential to fully exploit long and
    linked-read sequencing technologies.
acknowledgement: 'We thank the Barton group for useful discussion and feedback during
  the writing of this article. Comments from Roger Butlin, Molly Schumer''s Group,
  the tskit development team, editors and three reviewers greatly improved the manuscript.
  Funding was provided by SCAS (Natural Sciences Programme, Knut and Alice Wallenberg
  Foundation), an FWF Wittgenstein grant (PT1001Z211), an FWF standalone grant (grant
  P 32166), and an ERC Advanced Grant. YFC was supported by the Max Planck Society
  and an ERC Proof of Concept Grant #101069216 (HAPLOTAGGING).'
article_processing_charge: Yes (via OA deal)
article_type: original
author:
- first_name: Daria
  full_name: Shipilina, Daria
  id: 428A94B0-F248-11E8-B48F-1D18A9856A87
  last_name: Shipilina
  orcid: 0000-0002-1145-9226
- first_name: Arka
  full_name: Pal, Arka
  id: 6AAB2240-CA9A-11E9-9C1A-D9D1E5697425
  last_name: Pal
  orcid: 0000-0002-4530-8469
- first_name: Sean
  full_name: Stankowski, Sean
  id: 43161670-5719-11EA-8025-FABC3DDC885E
  last_name: Stankowski
- first_name: Yingguang Frank
  full_name: Chan, Yingguang Frank
  last_name: Chan
- first_name: Nicholas H
  full_name: Barton, Nicholas H
  id: 4880FE40-F248-11E8-B48F-1D18A9856A87
  last_name: Barton
  orcid: 0000-0002-8548-5240
citation:
  ama: Shipilina D, Pal A, Stankowski S, Chan YF, Barton NH. On the origin and structure
    of haplotype blocks. <i>Molecular Ecology</i>. 2023;32(6):1441-1457. doi:<a href="https://doi.org/10.1111/mec.16793">10.1111/mec.16793</a>
  apa: Shipilina, D., Pal, A., Stankowski, S., Chan, Y. F., &#38; Barton, N. H. (2023).
    On the origin and structure of haplotype blocks. <i>Molecular Ecology</i>. Wiley.
    <a href="https://doi.org/10.1111/mec.16793">https://doi.org/10.1111/mec.16793</a>
  chicago: Shipilina, Daria, Arka Pal, Sean Stankowski, Yingguang Frank Chan, and
    Nicholas H Barton. “On the Origin and Structure of Haplotype Blocks.” <i>Molecular
    Ecology</i>. Wiley, 2023. <a href="https://doi.org/10.1111/mec.16793">https://doi.org/10.1111/mec.16793</a>.
  ieee: D. Shipilina, A. Pal, S. Stankowski, Y. F. Chan, and N. H. Barton, “On the
    origin and structure of haplotype blocks,” <i>Molecular Ecology</i>, vol. 32,
    no. 6. Wiley, pp. 1441–1457, 2023.
  ista: Shipilina D, Pal A, Stankowski S, Chan YF, Barton NH. 2023. On the origin
    and structure of haplotype blocks. Molecular Ecology. 32(6), 1441–1457.
  mla: Shipilina, Daria, et al. “On the Origin and Structure of Haplotype Blocks.”
    <i>Molecular Ecology</i>, vol. 32, no. 6, Wiley, 2023, pp. 1441–57, doi:<a href="https://doi.org/10.1111/mec.16793">10.1111/mec.16793</a>.
  short: D. Shipilina, A. Pal, S. Stankowski, Y.F. Chan, N.H. Barton, Molecular Ecology
    32 (2023) 1441–1457.
corr_author: '1'
date_created: 2023-01-12T12:09:17Z
date_published: 2023-03-01T00:00:00Z
date_updated: 2026-05-05T22:31:12Z
day: '01'
ddc:
- '570'
department:
- _id: NiBa
doi: 10.1111/mec.16793
external_id:
  isi:
  - '000900762000001'
  pmid:
  - '36433653'
file:
- access_level: open_access
  checksum: b10e0f8fa3dc4d72aaf77a557200978a
  content_type: application/pdf
  creator: dernst
  date_created: 2023-08-16T08:15:41Z
  date_updated: 2023-08-16T08:15:41Z
  file_id: '14062'
  file_name: 2023_MolecularEcology_Shipilina.pdf
  file_size: 7144607
  relation: main_file
  success: 1
file_date_updated: 2023-08-16T08:15:41Z
has_accepted_license: '1'
intvolume: '        32'
isi: 1
issue: '6'
keyword:
- Genetics
- Ecology
- Evolution
- Behavior and Systematics
language:
- iso: eng
month: '03'
oa: 1
oa_version: Published Version
page: 1441-1457
pmid: 1
project:
- _id: 05959E1C-7A3F-11EA-A408-12923DDC885E
  grant_number: P32166
  name: Snapdragon Speciation
- _id: 25F42A32-B435-11E9-9278-68D0E5697425
  call_identifier: FWF
  grant_number: Z211
  name: Formal methods for the design and analysis of complex systems
- _id: bd6958e0-d553-11ed-ba76-86eba6a76c00
  grant_number: '101055327'
  name: Understanding the evolution of continuous genomes
publication: Molecular Ecology
publication_identifier:
  eissn:
  - 1365-294X
  issn:
  - 0962-1083
publication_status: published
publisher: Wiley
quality_controlled: '1'
related_material:
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  - id: '20694'
    relation: dissertation_contains
    status: public
scopus_import: '1'
status: public
title: On the origin and structure of haplotype blocks
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: journal_article
user_id: 2DF688A6-F248-11E8-B48F-1D18A9856A87
volume: 32
year: '2023'
...
---
_id: '1513'
abstract:
- lang: eng
  text: "Insects of the order Hemiptera (true bugs) use a wide range of mechanisms
    of sex determination, including genetic sex determination, paternal genome elimination,
    and haplodiploidy. Genetic sex determination, the prevalent mode, is generally
    controlled by a pair of XY sex chromosomes or by an XX/X0 system, but different
    configurations that include additional sex chromosomes are also present. Although
    this diversity of sex determining systems has been extensively studied at the
    cytogenetic level, only the X chromosome of the model pea aphid Acyrthosiphon
    pisum has been analyzed at the genomic level, and little is known about X chromosome
    biology in the rest of the order.\r\n\r\nIn this study, we take advantage of published
    DNA- and RNA-seq data from three additional Hemiptera species to perform a comparative
    analysis of the gene content and expression of the X chromosome throughout this
    clade. We find that, despite showing evidence of dosage compensation, the X chromosomes
    of these species show female-biased expression, and a deficit of male-biased genes,
    in direct contrast to the pea aphid X. We further detect an excess of shared gene
    content between these very distant species, suggesting that despite the diversity
    of sex determining systems, the same chromosomal element is used as the X throughout
    a large portion of the order. "
article_processing_charge: No
author:
- first_name: Arka
  full_name: Pal, Arka
  id: 6AAB2240-CA9A-11E9-9C1A-D9D1E5697425
  last_name: Pal
  orcid: 0000-0002-4530-8469
- first_name: Beatriz
  full_name: Vicoso, Beatriz
  id: 49E1C5C6-F248-11E8-B48F-1D18A9856A87
  last_name: Vicoso
  orcid: 0000-0002-4579-8306
citation:
  ama: 'Pal A, Vicoso B. The X chromosome of hemipteran insects: Conservation, dosage
    compensation and sex-biased expression. <i>Genome Biology and Evolution</i>. 2015;7(12):3259-3268.
    doi:<a href="https://doi.org/10.1093/gbe/evv215">10.1093/gbe/evv215</a>'
  apa: 'Pal, A., &#38; Vicoso, B. (2015). The X chromosome of hemipteran insects:
    Conservation, dosage compensation and sex-biased expression. <i>Genome Biology
    and Evolution</i>. Oxford University Press. <a href="https://doi.org/10.1093/gbe/evv215">https://doi.org/10.1093/gbe/evv215</a>'
  chicago: 'Pal, Arka, and Beatriz Vicoso. “The X Chromosome of Hemipteran Insects:
    Conservation, Dosage Compensation and Sex-Biased Expression.” <i>Genome Biology
    and Evolution</i>. Oxford University Press, 2015. <a href="https://doi.org/10.1093/gbe/evv215">https://doi.org/10.1093/gbe/evv215</a>.'
  ieee: 'A. Pal and B. Vicoso, “The X chromosome of hemipteran insects: Conservation,
    dosage compensation and sex-biased expression,” <i>Genome Biology and Evolution</i>,
    vol. 7, no. 12. Oxford University Press, pp. 3259–3268, 2015.'
  ista: 'Pal A, Vicoso B. 2015. The X chromosome of hemipteran insects: Conservation,
    dosage compensation and sex-biased expression. Genome Biology and Evolution. 7(12),
    3259–3268.'
  mla: 'Pal, Arka, and Beatriz Vicoso. “The X Chromosome of Hemipteran Insects: Conservation,
    Dosage Compensation and Sex-Biased Expression.” <i>Genome Biology and Evolution</i>,
    vol. 7, no. 12, Oxford University Press, 2015, pp. 3259–68, doi:<a href="https://doi.org/10.1093/gbe/evv215">10.1093/gbe/evv215</a>.'
  short: A. Pal, B. Vicoso, Genome Biology and Evolution 7 (2015) 3259–3268.
corr_author: '1'
date_created: 2018-12-11T11:52:27Z
date_published: 2015-12-01T00:00:00Z
date_updated: 2025-09-23T14:18:15Z
day: '01'
ddc:
- '570'
department:
- _id: BeVi
doi: 10.1093/gbe/evv215
ec_funded: 1
external_id:
  isi:
  - '000366498700008'
file:
- access_level: open_access
  checksum: 2b56b8c2e2a1d4cc3c9cb8daba26dd9b
  content_type: application/pdf
  creator: system
  date_created: 2018-12-12T10:17:29Z
  date_updated: 2020-07-14T12:45:00Z
  file_id: '5284'
  file_name: IST-2016-496-v1+1_Genome_Biol_Evol-2015-Pal-3259-68.pdf
  file_size: 858027
  relation: main_file
file_date_updated: 2020-07-14T12:45:00Z
has_accepted_license: '1'
intvolume: '         7'
isi: 1
issue: '12'
language:
- iso: eng
month: '12'
oa: 1
oa_version: Published Version
page: 3259 - 3268
project:
- _id: 25681D80-B435-11E9-9278-68D0E5697425
  call_identifier: FP7
  grant_number: '291734'
  name: International IST Postdoc Fellowship Programme
publication: Genome Biology and Evolution
publication_status: published
publisher: Oxford University Press
publist_id: '5664'
pubrep_id: '496'
quality_controlled: '1'
scopus_import: '1'
status: public
title: 'The X chromosome of hemipteran insects: Conservation, dosage compensation
  and sex-biased expression'
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: journal_article
user_id: 317138e5-6ab7-11ef-aa6d-ffef3953e345
volume: 7
year: '2015'
...
