{
  "abstract": "Host defense systems in prokaryotes and eukaryotes often appear to be derived from the very parasites they now defend against, yet the evolutionary route for such a transition remains poorly understood. Based on the biology of integrating viruses such as temperate bacteriophages and retroviruses, we develop a multilevel eco-evolutionary framework in which within-host viral replication is treated as a public goods game over a costly, shared replication product. In well-mixed host populations, we predict convergence on a single viral strategy of intermediate replicative investment. However, when we impose spatial structure by restricting host reproduction and virus transmission to local events, the virus population diversifies into two lineages: a horizontally transmitted lineage that parasitises hosts, and a vertically transmitted hyperparasitic lineage that parasitises the horizontally transmitting viruses. The latter \"hyperparasite\" is a viral cheat that suppresses the replication and propagation of the parasitic virus, thereby acting as a defense system to the hosts. Local interactions concentrate the benefits of this suppression among neighbouring hosts that carry related proviruses, allowing selection on the proviral cheat as a host-level defense. Our results show that evolutionary cheating within parasite populations can drive the evolution of parasites into defense systems in spatially structured host populations.",
  "abstract_shown": true,
  "affiliations": [
    "Utrecht University"
  ],
  "article_type": "New results",
  "arxiv": null,
  "author_total": 4,
  "authors": [
    {
      "affs": [
        0
      ],
      "name": "S. A. Nazir"
    },
    {
      "name": "B. Ashby"
    },
    {
      "name": "A. Leeks"
    },
    {
      "name": "B. van Dijk"
    }
  ],
  "bibtex": "https://research.draconitic.com/paper/2g0nk8tygqdxf.bib",
  "cited_by": 0,
  "doi": "10.64898/2026.10.14.759074",
  "fulltext_url": "https://www.biorxiv.org/content/10.64898/2026.10.14.759074v1.full",
  "id": "2g0nk8tygqdxf",
  "issns": [],
  "landing_url": "https://www.biorxiv.org/content/10.64898/2026.10.14.759074v1",
  "language": null,
  "licence": "CC-BY-NC",
  "licence_label": "CC BY-NC",
  "licence_source": "article",
  "licence_url": "https://creativecommons.org/licenses/by-nc/4.0/",
  "metadata_licence": null,
  "native_subjects": [
    [
      "biorxiv",
      "evolutionary biology"
    ],
    [
      "openalex",
      "1311|Evolution and Genetic Dynamics"
    ]
  ],
  "openalex": "W7222810179",
  "pdf_url": "https://www.biorxiv.org/content/10.64898/2026.10.14.759074v1.full.pdf",
  "pmcid": null,
  "pmid": null,
  "published": "2026-10-14",
  "publisher": null,
  "ris": "https://research.draconitic.com/paper/2g0nk8tygqdxf.ris",
  "sources": [
    {
      "first_seen": "2026-10-30T12:28:17Z",
      "native_id": "10.64898/2026.10.14.759074",
      "source": "biorxiv",
      "url": "https://www.biorxiv.org/content/10.64898/2026.10.14.759074v1"
    }
  ],
  "subjects": [
    {
      "field": "biology",
      "origin": "biorxiv:evolutionary biology",
      "subfield": "evolutionary-biology"
    }
  ],
  "title": "Spatial structure drives the evolution of parasites into host defense systems",
  "title_text": "Spatial structure drives the evolution of parasites into host defense systems",
  "type": "preprint",
  "updated_at": "2026-10-30T12:28:17Z",
  "url": "https://research.draconitic.com/paper/2g0nk8tygqdxf",
  "venue": "bioRxiv",
  "versions": []
}