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In host\u2010associated phytophagous insects, divergent host use can generate strong habitat isolation that evolves early in speciation and often results in sudden shifts towards strong reproductive isolation. However, whether additional sexual or postmating barriers can accumulate after this strong early ecological barrier has evolved remains unclear. In this study, we examined this question in the host\u2010specialised populations of the Southeast\u2010Asian herbivorous ladybird beetle,\n                    <jats:italic>Henosepilachna diekei<\/jats:italic>\n                    . We focused on the\n                    <jats:italic>Dicliptera\u2013Mikania<\/jats:italic>\n                    race pair and compared it with the previously studied\n                    <jats:italic>Mikania\u2013Leucas<\/jats:italic>\n                    race pair, representing two putative levels of host\u2010race differentiation. Combining laboratory crossing and field\u2010cage experiments, we investigated and quantified four potential isolating factors: premating isolation (habitat and sexual isolation), postmating\u2010prehatching isolation (reduced egg hatchability), and postmating\u2010posthatching isolation (F\n                    <jats:sub>1<\/jats:sub>\n                    hybrid inviability). The\n                    <jats:italic>Dicliptera<\/jats:italic>\n                    \u2013\n                    <jats:italic>Mikania<\/jats:italic>\n                    race pair showed larger genetic differentiation than the sympatric\n                    <jats:italic>Mikania<\/jats:italic>\n                    \u2013\n                    <jats:italic>Leucas<\/jats:italic>\n                    race pair. Both race pairs showed strong habitat isolation, but only the\n                    <jats:italic>Dicliptera<\/jats:italic>\n                    \u2013\n                    <jats:italic>Mikania<\/jats:italic>\n                    race pair showed additional later\u2010acting barriers, including reduced mating success, reduced egg hatchability, and reduced F\n                    <jats:sub>1<\/jats:sub>\n                    hybrid survival. Our findings suggest that additional reproductive barriers can evolve even in the presence of exceptionally strong habitat isolation during host\u2010race divergence. Rather than replacing the importance of the initial host\u2010associated barrier, late\u2010acting barriers may accumulate and contribute to the stability of reproductive isolation. Consequently, the strength and dynamics of later\u2010acting barriers may depend on the evolutionary trajectory of each host\u2010race pair, reflecting diverse pathways through which speciation can unfold in host\u2010associated insects.\n                  <\/jats:p>","DOI":"10.1002\/ece3.74046","type":"journal-article","created":{"date-parts":[[2026,7,17]],"date-time":"2026-07-17T13:56:19Z","timestamp":1784296579000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["The Evolution of Reproductive Isolation Beyond a Strong First Barrier in Speciation Between Micro\u2010Allopatric Host Races of a Phytophagous Ladybird Beetle,\n                    <i>Henosepilachna diekei<\/i>"],"prefix":"10.1002","volume":"16","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1968-4575","authenticated-orcid":false,"given":"Arif","family":"Maulana","sequence":"first","affiliation":[{"name":"Department of Biology, Faculty of Mathematics and Natural Sciences IPB University (Bogor Agricultural University)  Bogor West Java Indonesia"},{"name":"Wellcome Sanger Institute, Tree of Life Programme  Cambridge UK"},{"name":"Darwin College University of Cambridge  Cambridge UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1157-5622","authenticated-orcid":false,"given":"Kei W.","family":"Matsubayashi","sequence":"additional","affiliation":[{"name":"Department of Environmental Sciences, College of Agriculture, Food and Environment Sciences Rakuno Gakuen University  Ebetsu\u2010shi Hokkaido Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1483-4471","authenticated-orcid":false,"given":"Naoyuki","family":"Fujiyama","sequence":"additional","affiliation":[{"name":"Department of Biology, Faculty of Science Yamagata University  Yamagata Yamagata Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2179-3914","authenticated-orcid":false,"given":"Tri","family":"Atmowidi","sequence":"additional","affiliation":[{"name":"Department of Biology, Faculty of Mathematics and Natural Sciences IPB University (Bogor Agricultural University)  Bogor West Java Indonesia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7222-3180","authenticated-orcid":false,"given":"Sih","family":"Kahono","sequence":"additional","affiliation":[{"name":"National Research and Innovation Agency Research Centre for Biota Systems, Kawasan Sains dan Teknologi Dr. (H.C.) 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