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(PDF) Quantitative genetic inheritance of morphological divergence in a lake-stream stickleback ecotype pair: implications for reproductive isolation | Andrew Hendry - Academia.edu

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{"work":{"id":21695199,"created_at":"2016-02-08T09:18:47.077-08:00","from_world_paper_id":148163983,"updated_at":"2021-01-13T01:55:51.957-08:00","_data":{"publication_date":"2011,,","publication_name":"Journal of Evolutionary Biology"},"document_type":"paper","pre_hit_view_count_baseline":null,"quality":"high","language":"en","title":"Quantitative genetic inheritance of morphological divergence in a lake-stream stickleback ecotype pair: implications for reproductive isolation","broadcastable":false,"draft":null,"has_indexable_attachment":true,"indexable":true}}["work"]; window.loswp.workCoauthors = [32975920]; window.loswp.locale = "en"; window.loswp.countryCode = "SG"; window.loswp.cwvAbTestBucket = ""; window.loswp.designVariant = "ds_vanilla"; window.loswp.fullPageMobileSutdModalVariant = "full_page_mobile_sutd_modal"; window.loswp.useOptimizedScribd4genScript = false; window.loswp.appleClientId = 'edu.academia.applesignon';</script><script defer="" 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src="//a.academia-assets.com/images/s65_no_pic.png" />Andrew Hendry</a></div><div class="ds-work-card--detail"><p class="ds-work-card--detail ds2-5-body-sm">2011, Journal of Evolutionary Biology</p></div><div class="ds-work-card--button-container"><button class="ds2-5-button js-swp-download-button" data-signup-modal="{&quot;location&quot;:&quot;continue-reading-button--work-card&quot;,&quot;attachmentId&quot;:42393157,&quot;attachmentType&quot;:&quot;pdf&quot;,&quot;workUrl&quot;:&quot;https://www.academia.edu/21695199/Quantitative_genetic_inheritance_of_morphological_divergence_in_a_lake_stream_stickleback_ecotype_pair_implications_for_reproductive_isolation&quot;}">See full PDF</button><button class="ds2-5-button ds2-5-button--secondary js-swp-download-button" 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ds2-5-body-sm ds2-5-body-link" data-author-id="4533491" href="https://ubc.academia.edu/JenniferGow">Jennifer Gow</a></div><p class="ds-related-work--metadata ds2-5-body-xs">2007</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{&quot;location&quot;:&quot;wsj-grid-card-download-pdf-modal&quot;,&quot;work_title&quot;:&quot;Quantifying the constraining influence of gene flow on adaptive divergence in the lake-stream threespine stickleback system&quot;,&quot;attachmentId&quot;:31694997,&quot;attachmentType&quot;:&quot;pdf&quot;,&quot;work_url&quot;:&quot;https://www.academia.edu/4200004/Quantifying_the_constraining_influence_of_gene_flow_on_adaptive_divergence_in_the_lake_stream_threespine_stickleback_system&quot;,&quot;alternativeTracking&quot;:true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free 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class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="32975920" href="https://independent.academia.edu/AndrewHendry">Andrew Hendry</a></div><p class="ds-related-work--metadata ds2-5-body-xs">2012</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{&quot;location&quot;:&quot;wsj-grid-card-download-pdf-modal&quot;,&quot;work_title&quot;:&quot;Divergent Selection and Then What Not: The Conundrum of Missing Reproductive Isolation in Misty Lake and Stream Stickleback&quot;,&quot;attachmentId&quot;:42393105,&quot;attachmentType&quot;:&quot;pdf&quot;,&quot;work_url&quot;:&quot;https://www.academia.edu/21695131/Divergent_Selection_and_Then_What_Not_The_Conundrum_of_Missing_Reproductive_Isolation_in_Misty_Lake_and_Stream_Stickleback&quot;,&quot;alternativeTracking&quot;:true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/21695131/Divergent_Selection_and_Then_What_Not_The_Conundrum_of_Missing_Reproductive_Isolation_in_Misty_Lake_and_Stream_Stickleback"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="8" data-entity-id="25210595" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/25210595/Where_the_Lake_Meets_the_Sea_Strong_Reproductive_Isolation_Is_Associated_with_Adaptive_Divergence_between_Lake_Resident_and_Anadromous_Three_Spined_Sticklebacks">Where the Lake Meets the Sea: Strong Reproductive Isolation Is Associated with Adaptive Divergence between Lake Resident and Anadromous Three-Spined Sticklebacks</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="4842" href="https://glasgow.academia.edu/ChrisHarrod">Chris Harrod</a></div><p class="ds-related-work--metadata ds2-5-body-xs">PloS one, 2015</p><p class="ds-related-work--abstract ds2-5-body-sm">Contact zones between divergent forms of the same species are often characterised by high levels of phenotypic diversity over small geographic distances. What processes are involved in generating such high phenotypic diversity? One possibility is that introgression and recombination between divergent forms in contact zones results in greater phenotypic and genetic polymorphism. Alternatively, strong reproductive isolation between forms may maintain distinct phenotypes, preventing homogenisation by gene flow. Contact zones between divergent freshwater-resident and anadromous stickleback (Gasterosteus aculeatus L.) forms are numerous and common throughout the species distribution, offering an opportunity to examine these contrasting hypotheses in greater detail. This study reports on an interesting new contact zone located in a tidally influenced lake catchment in western Ireland, characterised by high polymorphism for lateral plate phenotypes. Using neutral and QTL-linked microsatell...</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{&quot;location&quot;:&quot;wsj-grid-card-download-pdf-modal&quot;,&quot;work_title&quot;:&quot;Where the Lake Meets the Sea: Strong Reproductive Isolation Is Associated with Adaptive Divergence between Lake Resident and Anadromous Three-Spined Sticklebacks&quot;,&quot;attachmentId&quot;:45521469,&quot;attachmentType&quot;:&quot;pdf&quot;,&quot;work_url&quot;:&quot;https://www.academia.edu/25210595/Where_the_Lake_Meets_the_Sea_Strong_Reproductive_Isolation_Is_Associated_with_Adaptive_Divergence_between_Lake_Resident_and_Anadromous_Three_Spined_Sticklebacks&quot;,&quot;alternativeTracking&quot;:true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/25210595/Where_the_Lake_Meets_the_Sea_Strong_Reproductive_Isolation_Is_Associated_with_Adaptive_Divergence_between_Lake_Resident_and_Anadromous_Three_Spined_Sticklebacks"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-wsj-grid-card" data-collection-position="9" data-entity-id="3704910" data-sort-order="default"><a class="ds-related-work--title js-wsj-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/3704910/Speciation_in_reverse_morphological_and_genetic_evidence_of_the_collapse_of_a_three_spined_stickleback_Gasterosteus_aculeatus_species_pair">Speciation in reverse: morphological and genetic evidence of the collapse of a three-spined stickleback (Gasterosteus aculeatus) species pair</a><div class="ds-related-work--metadata"><a class="js-wsj-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="4533491" href="https://ubc.academia.edu/JenniferGow">Jennifer Gow</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Molecular Ecology, 2005</p><p class="ds-related-work--abstract ds2-5-body-sm">Historically, six small lakes in southwestern British Columbia each contained a sympatric species pair of three-spined sticklebacks (Gasterosteus aculeatus). These pairs consisted of a ‘benthic’ and ‘limnetic’ species that had arisen postglacially and, in four of the lakes, independently. Sympatric sticklebacks are considered biological species because they are morphologically, ecologically and genetically distinct and because they are strongly reproductively isolated from one another. The restricted range of the species pairs places them at risk of extinction, and one of the pairs has gone extinct after the introduction of an exotic catfish. In another lake, Enos Lake, southeastern Vancouver Island, an earlier report suggested that its species pair is at risk from elevated levels of hybridization. We conducted a detailed morphological analysis, as well as genetic analysis of variation at five microsatellite loci for samples spanning a time frame of 1977 to 2002 to test the hypothesis that the pair in Enos Lake is collapsing into a hybrid swarm. Our morphological analysis showed a clear breakdown between benthics and limnetics. Bayesian model-based clustering indicated that two morphological clusters were evident in 1977 and 1988, which were replaced by 1997 by a single highly variable cluster. The most recent 2000 and 2002 samples confirm the breakdown. Microsatellite analysis corroborated the morphological results. Bayesian analyses of population structure in a sample collected in 1994 indicated two genetically distinct populations in Enos Lake, but only a single genetic population was evident in 1997, 2000, and 2002. In addition, genetic analyses of samples collected in 1997, 2000, and 2002 showed strong signals of ‘hybrids’; they were genetically intermediate to parental genotypes. Our results support the idea that the Enos Lake species pair is collapsing into a hybrid swarm. Although the precise mechanism(s) responsible for elevated hybridization in the lake is unknown, the demise of the Enos Lake species pair follows the appearance of an exotic crayfish, Pascifasticus lenisculus, in the early 1990s.</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{&quot;location&quot;:&quot;wsj-grid-card-download-pdf-modal&quot;,&quot;work_title&quot;:&quot;Speciation in reverse: morphological and genetic evidence of the collapse of a three-spined stickleback (Gasterosteus aculeatus) species pair&quot;,&quot;attachmentId&quot;:31695051,&quot;attachmentType&quot;:&quot;pdf&quot;,&quot;work_url&quot;:&quot;https://www.academia.edu/3704910/Speciation_in_reverse_morphological_and_genetic_evidence_of_the_collapse_of_a_three_spined_stickleback_Gasterosteus_aculeatus_species_pair&quot;,&quot;alternativeTracking&quot;:true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-wsj-grid-card-view-pdf" href="https://www.academia.edu/3704910/Speciation_in_reverse_morphological_and_genetic_evidence_of_the_collapse_of_a_three_spined_stickleback_Gasterosteus_aculeatus_species_pair"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div></div></div><div class="ds-sticky-ctas--wrapper js-loswp-sticky-ctas hidden"><div class="ds-sticky-ctas--grid-container"><div class="ds-sticky-ctas--container"><button class="ds2-5-button js-swp-download-button" data-signup-modal="{&quot;location&quot;:&quot;continue-reading-button--sticky-ctas&quot;,&quot;attachmentId&quot;:42393157,&quot;attachmentType&quot;:&quot;pdf&quot;,&quot;workUrl&quot;:null}">See full PDF</button><button class="ds2-5-button ds2-5-button--secondary js-swp-download-button" data-signup-modal="{&quot;location&quot;:&quot;download-pdf-button--sticky-ctas&quot;,&quot;attachmentId&quot;:42393157,&quot;attachmentType&quot;:&quot;pdf&quot;,&quot;workUrl&quot;:null}"><span class="material-symbols-outlined" style="font-size: 20px" translate="no">download</span>Download PDF</button></div></div></div><div class="ds-below-fold--grid-container"><div class="ds-work--container js-loswp-embedded-document"><div class="attachment_preview" data-attachment="Attachment_42393157" style="display: none"><div class="js-scribd-document-container"><div class="scribd--document-loading js-scribd-document-loader" style="display: block;"><img alt="Loading..." src="//a.academia-assets.com/images/loaders/paper-load.gif" /><p>Loading Preview</p></div></div><div style="text-align: center;"><div class="scribd--no-preview-alert js-preview-unavailable"><p>Sorry, preview is currently unavailable. You can download the paper by clicking the button above.</p></div></div></div></div><div class="ds-sidebar--container js-work-sidebar"><div class="ds-related-content--container"><h2 class="ds-related-content--heading">Related papers</h2><div class="ds-related-work--container js-related-work-sidebar-card" data-collection-position="0" data-entity-id="12053090" data-sort-order="default"><a class="ds-related-work--title js-related-work-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/12053090/Natural_selection_drives_patterns_of_lake_stream_divergence_in_stickleback_foraging_morphology">Natural selection drives patterns of lake–stream divergence in stickleback foraging morphology</a><div class="ds-related-work--metadata"><a class="js-related-work-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="30047083" href="https://independent.academia.edu/DeanAdams3">Dean Adams</a></div><p class="ds-related-work--metadata ds2-5-body-xs">2008</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{&quot;location&quot;:&quot;wsj-grid-card-download-pdf-modal&quot;,&quot;work_title&quot;:&quot;Natural selection drives patterns of lake–stream divergence in stickleback foraging morphology&quot;,&quot;attachmentId&quot;:46392404,&quot;attachmentType&quot;:&quot;pdf&quot;,&quot;work_url&quot;:&quot;https://www.academia.edu/12053090/Natural_selection_drives_patterns_of_lake_stream_divergence_in_stickleback_foraging_morphology&quot;,&quot;alternativeTracking&quot;:true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-related-work-grid-card-view-pdf" href="https://www.academia.edu/12053090/Natural_selection_drives_patterns_of_lake_stream_divergence_in_stickleback_foraging_morphology"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-related-work-sidebar-card" data-collection-position="1" data-entity-id="25550529" data-sort-order="default"><a class="ds-related-work--title js-related-work-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/25550529/Taylor_EB_Boughman_JW_Groenenboom_M_Sniatynski_M_Schluter_D_Gow_JL_Speciation_in_reverse_morphological_and_genetic_evidence_of_the_collape_of_a_three_spined_stickleback_Gasterosteus_aculeatus_species_pair_Mol_Ecol_15_343_355">Taylor EB, Boughman JW, Groenenboom M, Sniatynski M, Schluter D, Gow JL. Speciation in reverse: morphological and genetic evidence of the collape of a three-spined stickleback (Gasterosteus aculeatus) species pair. Mol Ecol 15: 343-355</a><div class="ds-related-work--metadata"><a class="js-related-work-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="49124796" href="https://independent.academia.edu/JanetteBoughman">Janette Boughman</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Molecular Ecology</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{&quot;location&quot;:&quot;wsj-grid-card-download-pdf-modal&quot;,&quot;work_title&quot;:&quot;Taylor EB, Boughman JW, Groenenboom M, Sniatynski M, Schluter D, Gow JL. Speciation in reverse: morphological and genetic evidence of the collape of a three-spined stickleback (Gasterosteus aculeatus) species pair. Mol Ecol 15: 343-355&quot;,&quot;attachmentId&quot;:45882279,&quot;attachmentType&quot;:&quot;pdf&quot;,&quot;work_url&quot;:&quot;https://www.academia.edu/25550529/Taylor_EB_Boughman_JW_Groenenboom_M_Sniatynski_M_Schluter_D_Gow_JL_Speciation_in_reverse_morphological_and_genetic_evidence_of_the_collape_of_a_three_spined_stickleback_Gasterosteus_aculeatus_species_pair_Mol_Ecol_15_343_355&quot;,&quot;alternativeTracking&quot;:true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-related-work-grid-card-view-pdf" href="https://www.academia.edu/25550529/Taylor_EB_Boughman_JW_Groenenboom_M_Sniatynski_M_Schluter_D_Gow_JL_Speciation_in_reverse_morphological_and_genetic_evidence_of_the_collape_of_a_three_spined_stickleback_Gasterosteus_aculeatus_species_pair_Mol_Ecol_15_343_355"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-related-work-sidebar-card" data-collection-position="2" data-entity-id="21695249" data-sort-order="default"><a class="ds-related-work--title js-related-work-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/21695249/Asymmetric_reproductive_barriers_and_mosaic_reproductive_isolation_insights_from_Misty_lake_stream_stickleback">Asymmetric reproductive barriers and mosaic reproductive isolation: insights from Misty lake-stream stickleback</a><div class="ds-related-work--metadata"><a class="js-related-work-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="32975920" href="https://independent.academia.edu/AndrewHendry">Andrew Hendry</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Ecology and Evolution, 2014</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link 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class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-related-work-sidebar-card" data-collection-position="3" data-entity-id="3704912" data-sort-order="default"><a class="ds-related-work--title js-related-work-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/3704912/Contrasting_hybridization_rates_between_sympatric_three_spined_sticklebacks_highlight_the_fragility_of_reproductive_barriers_between_evolutionarily_young_species">Contrasting hybridization rates between sympatric three-spined sticklebacks highlight the fragility of reproductive barriers between evolutionarily young species</a><div class="ds-related-work--metadata"><a class="js-related-work-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="4533491" href="https://ubc.academia.edu/JenniferGow">Jennifer Gow</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Molecular Ecology, 2006</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{&quot;location&quot;:&quot;wsj-grid-card-download-pdf-modal&quot;,&quot;work_title&quot;:&quot;Contrasting hybridization rates between sympatric three-spined sticklebacks highlight the fragility of reproductive barriers between evolutionarily young species&quot;,&quot;attachmentId&quot;:31694461,&quot;attachmentType&quot;:&quot;pdf&quot;,&quot;work_url&quot;:&quot;https://www.academia.edu/3704912/Contrasting_hybridization_rates_between_sympatric_three_spined_sticklebacks_highlight_the_fragility_of_reproductive_barriers_between_evolutionarily_young_species&quot;,&quot;alternativeTracking&quot;:true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span class="ds2-5-text-link__content">Download free PDF</span></button><a class="ds2-5-text-link ds2-5-text-link--inline js-related-work-grid-card-view-pdf" href="https://www.academia.edu/3704912/Contrasting_hybridization_rates_between_sympatric_three_spined_sticklebacks_highlight_the_fragility_of_reproductive_barriers_between_evolutionarily_young_species"><span class="ds2-5-text-link__content">View PDF</span><span class="material-symbols-outlined" style="font-size: 18px" translate="no">chevron_right</span></a></div></div><div class="ds-related-work--container js-related-work-sidebar-card" data-collection-position="4" data-entity-id="115204048" data-sort-order="default"><a class="ds-related-work--title js-related-work-grid-card-title ds2-5-body-md ds2-5-body-link" href="https://www.academia.edu/115204048/Adaptive_Divergence_under_Gene_Flow_along_an_Environmental_Gradient_in_Two_Coexisting_Stickleback_Species">Adaptive Divergence under Gene Flow along an Environmental Gradient in Two Coexisting Stickleback Species</a><div class="ds-related-work--metadata"><a class="js-related-work-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="221914350" href="https://independent.academia.edu/BHellemans">Bart Hellemans</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Genes, 2021</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{&quot;location&quot;:&quot;wsj-grid-card-download-pdf-modal&quot;,&quot;work_title&quot;:&quot;Adaptive Divergence under Gene Flow along an Environmental Gradient in Two Coexisting Stickleback Species&quot;,&quot;attachmentId&quot;:111680003,&quot;attachmentType&quot;:&quot;pdf&quot;,&quot;work_url&quot;:&quot;https://www.academia.edu/115204048/Adaptive_Divergence_under_Gene_Flow_along_an_Environmental_Gradient_in_Two_Coexisting_Stickleback_Species&quot;,&quot;alternativeTracking&quot;:true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span 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class="js-related-work-grid-card-author ds2-5-body-sm ds2-5-body-link" data-author-id="7541539" href="https://independent.academia.edu/oleseehausen">ole seehausen</a></div><p class="ds-related-work--metadata ds2-5-body-xs">Nature Ecology and Evolution, 2017</p><div class="ds-related-work--ctas"><button class="ds2-5-text-link ds2-5-text-link--inline js-swp-download-button" data-signup-modal="{&quot;location&quot;:&quot;wsj-grid-card-download-pdf-modal&quot;,&quot;work_title&quot;:&quot;Transgenerational selection driven by divergent ecological impacts of hybridizing lineages&quot;,&quot;attachmentId&quot;:111785612,&quot;attachmentType&quot;:&quot;pdf&quot;,&quot;work_url&quot;:&quot;https://www.academia.edu/115353827/Transgenerational_selection_driven_by_divergent_ecological_impacts_of_hybridizing_lineages&quot;,&quot;alternativeTracking&quot;:true}"><span class="material-symbols-outlined" style="font-size: 18px" translate="no">download</span><span 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