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Five years of integrated surveillance (2020\u20132024) at three Maryland sites allowed us to track\n                      <jats:italic toggle=\"yes\">B. microti<\/jats:italic>\n                      and\n                      <jats:italic toggle=\"yes\">A. phagocytophilum<\/jats:italic>\n                      establishment by screening questing\n                      <jats:italic toggle=\"yes\">Ixodes scapularis<\/jats:italic>\n                      nymphs,\n                      <jats:italic toggle=\"yes\">Peromyscus leucopus-<\/jats:italic>\n                      fed nymphs, and\n                      <jats:italic toggle=\"yes\">P. leucopus<\/jats:italic>\n                      samples by qPCR, contextualized with county-level human case data.\n                      <jats:italic toggle=\"yes\">B. burgdorferi<\/jats:italic>\n                      was consistently detected in all sites and sample types, with prevalence generally 5%\u201320% in questing nymphs and exceeding 30% in hosts, confirming long-term enzootic maintenance. By contrast,\n                      <jats:italic toggle=\"yes\">B. microti<\/jats:italic>\n                      and\n                      <jats:italic toggle=\"yes\">A. phagocytophilum<\/jats:italic>\n                      were initially sporadic but increased in rodents and\n                      <jats:italic toggle=\"yes\">P. leucopus-<\/jats:italic>\n                      fed ticks. Over time,\n                      <jats:italic toggle=\"yes\">A. phagocytophilum<\/jats:italic>\n                      prevalence significantly increased to above 20% in some\n                      <jats:italic toggle=\"yes\">P. leucopus-<\/jats:italic>\n                      fed nymphal collections despite much lower prevalence in questing ticks, highlighting the early-warning value of blood meal-associated surveillance. Notably,\n                      <jats:italic toggle=\"yes\">B. microti<\/jats:italic>\n                      reached very high prevalence in\n                      <jats:italic toggle=\"yes\">P. leucopus<\/jats:italic>\n                      hosts at a specific site (up to ~80%) while remaining rare or absent in questing and engorged nymphs, highlighting a pronounced decoupling between host infection and vector prevalence. Coinfections were rare, though enrichment of\n                      <jats:italic toggle=\"yes\">B. burgdorferi + A. phagocytophilum<\/jats:italic>\n                      in\n                      <jats:italic toggle=\"yes\">P. leucopus-<\/jats:italic>\n                      fed ticks suggests possible facilitation during early establishment. These results indicate that\n                      <jats:italic toggle=\"yes\">B. microti<\/jats:italic>\n                      and\n                      <jats:italic toggle=\"yes\">A. phagocytophilum<\/jats:italic>\n                      are actively emerging in Maryland, following their establishment in the Northeast and Upper Midwest. Combining surveillance from questing nymphs,\n                      <jats:italic toggle=\"yes\">P. leucopus-<\/jats:italic>\n                      fed nymphs, and reservoir hosts provides a framework for detecting enzootic cycles before they appear in questing ticks or human case counts, offering early-warning capacity for public health preparedness.\n                    <\/jats:p>\n                    <jats:sec>\n                      <jats:title>IMPORTANCE<\/jats:title>\n                      <jats:p>\n                        Understanding why some tick-borne pathogens become ecologically established while others remain sporadic is central to predicting human disease emergence. By combining surveillance of questing nymphal\n                        <jats:italic toggle=\"yes\">Ixodes scapularis<\/jats:italic>\n                        ticks,\n                        <jats:italic toggle=\"yes\">Peromyscus leucopus-<\/jats:italic>\n                        fed nymphal ticks, and\n                        <jats:italic toggle=\"yes\">Peromyscus leucopus<\/jats:italic>\n                        reservoir hosts across 5 years in Maryland, we show that\n                        <jats:italic toggle=\"yes\">Babesia microti<\/jats:italic>\n                        and\n                        <jats:italic toggle=\"yes\">Anaplasma phagocytophilum<\/jats:italic>\n                        remain in the early stages of enzootic establishment, whereas\n                        <jats:italic toggle=\"yes\">Borrelia burgdorferi<\/jats:italic>\n                        is deeply established. This integrated approach demonstrates how pathogen biology within the tick shapes field prevalence and highlights\n                        <jats:italic toggle=\"yes\">P. leucopus-<\/jats:italic>\n                        fed ticks as a powerful xenodiagnostic early-warning tool for detecting emerging pathogens before they are reflected in questing populations or human case data.\n                      <\/jats:p>\n                    <\/jats:sec>\n                  <\/jats:sec>","DOI":"10.1128\/aem.02242-25","type":"journal-article","created":{"date-parts":[[2026,4,20]],"date-time":"2026-04-20T13:00:44Z","timestamp":1776690044000},"update-policy":"https:\/\/doi.org\/10.1128\/asmj-crossmark-policy-page","source":"Crossref","is-referenced-by-count":0,"title":["Coinfection ecology and pathogen emergence in a\n                    <i>Borrelia<\/i>\n                    -endemic landscape: 5 years of\n                    <i>Borrelia burgdorferi<\/i>\n                    ,\n                    <i>Anaplasma phagocytophilum<\/i>\n                    , and\n                    <i>Babesia microti<\/i>\n                    surveillance in Maryland"],"prefix":"10.1128","volume":"92","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9333-6600","authenticated-orcid":false,"given":"Greg","family":"Joyner","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0011qv509","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Microbiology, Immunology and Biochemistry, University of Tennessee Health Science Center","place":["Memphis, USA"]},{"id":[{"id":"https:\/\/ror.org\/016ata990","id-type":"ROR","asserted-by":"publisher"}],"name":"Immuno Technologies Inc.","place":["Memphis, USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Olifan","family":"Abil","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0011qv509","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Microbiology, Immunology and Biochemistry, 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