Spatially limited pathogen pollution in an invasive tick and host system

<!DOCTYPE html> <html lang="en" > <head > <meta charset="UTF-8" /> <meta http-equiv="X-UA-Compatible" content="IE=edge" /> <meta name="HandheldFriendly" content="True" /> <meta name="MobileOptimized" content="320" /> <meta name="viewport" content="width=device-width, initial-scale=1.0" /> <link rel="stylesheet" href="/static/assets/style-70b9163a.css" /> <script type="module" crossorigin="" src="/static/assets/base_style-ec2bc71e.js"></script> <link rel="stylesheet" href="/static/assets/style-ef962842.css" /> <link rel="stylesheet" href="/static/assets/style-3ade8b5c.css" /> <script type="module" crossorigin="" src="/static/assets/article_style-d757a0dd.js"></script> <style> @media screen and (min-width: 64em) { div.pmc-wm { background: repeat-y; background-image: url("data:image/svg+xml,%3Csvg xmlns='http://www.w3.org/2000/svg' width='20' height='350' xmlns:xlink='http://www.w3.org/1999/xlink'%3E%3Cdefs%3E%3Cfilter x='-.02' y='0' width='1.05' height='1' id='c'%3E%3CfeFlood flood-color='%23FFF'/%3E%3CfeComposite in='SourceGraphic'/%3E%3C/filter%3E%3Ctext id='b' font-family='Helvetica' font-size='11pt' style='opacity:1;fill:%23005ea2;stroke:none;text-anchor:middle' x='175' y='14'%3E%3C/text%3E%3Cpath id='a' style='fill:%23005ea2' d='M0 8h350v3H0z'/%3E%3C/defs%3E%3Cuse xlink:href='%23a' transform='rotate(90 10 10)'/%3E%3Cuse xlink:href='%23b' transform='rotate(90 10 10)' filter='url(%23c)'/%3E%3C/svg%3E"); padding-left: 3rem; } } </style> <link rel="apple-touch-icon" sizes="180x180" href="/static/img/favicons/apple-touch-icon.png" /> <link rel="icon" type="image/png" sizes="48x48" href="/static/img/favicons/favicon-48x48.png" /> <link rel="icon" type="image/png" sizes="32x32" href="/static/img/favicons/favicon-32x32.png" /> <link rel="icon" type="image/png" sizes="16x16" href="/static/img/favicons/favicon-16x16.png" /> <link rel="manifest" href="/static/img/favicons/site.webmanifest" /> <link rel="mask-icon" href="/static/img/favicons/safari-pinned-tab.svg" color="#0071bc" /> <meta name="msapplication-config" content="/static/img/favicons/browserconfig.xml" /> <meta name="theme-color" content="#ffffff" /> <title> Spatially limited pathogen pollution in an invasive tick and host system - PMC </title>  <meta name="ncbi_app" content="cloudpmc-viewer" /> <meta name="ncbi_db" content="pmc" /> <meta name="ncbi_phid" content="76C100057465D733060005000C1CCAC7.m_1" />  <meta name="ncbi_pdid" content="article" /> <link rel="preconnect" href="https://www.google-analytics.com" /> <link rel="dns-prefetch" href="https://cdn.ncbi.nlm.nih.gov" /> <link rel="preconnect" href="https://code.jquery.com" /> <meta name="ncbi_domain" content="springeropen"> <meta name="ncbi_type" content="fulltext"> <meta name="ncbi_pcid" content="journal"> <meta name="ncbi_feature" content="associated_data"> <link rel="canonical" href="https://pmc.ncbi.nlm.nih.gov/articles/PMC11213802/"> <meta name="robots" content="INDEX,NOFOLLOW,NOARCHIVE"> <meta name="citation_journal_title" content="Biological Invasions"> <meta name="citation_title" content="Spatially limited pathogen pollution in an invasive tick and host system"> <meta name="citation_author" content="Carrie E De Jesus"> <meta name="citation_author_institution" content="Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, FL USA"> <meta name="citation_author" content="Madison E A Harman"> <meta name="citation_author_institution" content="Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, FL USA"> <meta name="citation_author" content="Amber Sutton"> <meta name="citation_author_institution" content="Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, FL USA"> <meta name="citation_author" content="Stephen Bredin"> <meta name="citation_author_institution" content="Biology Department, George Mason University, Fairfax, VA USA"> <meta name="citation_author" content="Christina M Romagosa"> <meta name="citation_author_institution" content="Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, FL USA"> <meta name="citation_author" content="Samantha M Wisely"> <meta name="citation_author_institution" content="Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, FL USA"> <meta name="citation_publication_date" content="2024 Apr 23"> <meta name="citation_volume" content="26"> <meta name="citation_issue" content="7"> <meta name="citation_firstpage" content="2037"> <meta name="citation_doi" content="10.1007/s10530-024-03291-9"> <meta name="citation_pmid" content="38947421"> <meta name="citation_abstract_html_url" content="https://pmc.ncbi.nlm.nih.gov/articles/PMC11213802/"> <meta name="citation_fulltext_html_url" content="https://pmc.ncbi.nlm.nih.gov/articles/PMC11213802/"> <meta name="citation_pdf_url" content="https://pmc.ncbi.nlm.nih.gov/articles/PMC11213802/pdf/10530_2024_Article_3291.pdf"> <meta name="description" content="Expansion of global commerce has facilitated pathogen pollution via the transportation and translocation of invasive species and their associated parasites and pathogens. In Florida, imported cane toads (Rhinella horribilis) were accidentally and ..."> <meta name="og:title" content="Spatially limited pathogen pollution in an invasive tick and host system"> <meta name="og:type" content="article"> <meta name="og:site_name" content="PubMed Central (PMC)"> <meta name="og:description" content="Expansion of global commerce has facilitated pathogen pollution via the transportation and translocation of invasive species and their associated parasites and pathogens. In Florida, imported cane toads (Rhinella horribilis) were accidentally and ..."> <meta name="og:url" content="https://pmc.ncbi.nlm.nih.gov/articles/PMC11213802/"> <meta name="og:image" content="https://cdn.ncbi.nlm.nih.gov/pmc/cms/images/pmc-card-share.jpg?_=0"> <meta name="twitter:card" content="summary_large_image"> <meta name="twitter:site" content="@ncbi"> </head> <body > <a class="usa-skipnav " href="#main-content"> Skip to main content </a> <section class="usa-banner " aria-label="Official website of the United States government" > <div class="usa-accordion"> <header class="usa-banner__header"> <div class="usa-banner__inner"> <div class="grid-col-auto"> <img aria-hidden="true" class="usa-banner__header-flag" src="/static/img/us_flag.svg" alt="" /> </div> <div class="grid-col-fill tablet:grid-col-auto" aria-hidden="true"> <p class="usa-banner__header-text"> An official website of the United States government </p> <span class="usa-banner__header-action">Here's how you know</span> </div> <button type="button" class="usa-accordion__button usa-banner__button " aria-expanded="false" aria-controls="gov-banner-default" data-testid="storybook-django-banner" > <span class="usa-banner__button-text">Here's how you know</span> </button> </div> </header> <div class="usa-banner__content usa-accordion__content" id="gov-banner-default" hidden> <div class="grid-row grid-gap-lg"> <div class="usa-banner__guidance tablet:grid-col-6"> <img class="usa-banner__icon usa-media-block__img" src="/static/img/icon-dot-gov.svg" alt="" aria-hidden="true" /> <div class="usa-media-block__body"> <p> <strong>Official websites use .gov</strong> <br /> A <strong>.gov</strong> website belongs to an official government organization in the United States. </p> </div> </div> <div class="usa-banner__guidance tablet:grid-col-6"> <img class="usa-banner__icon usa-media-block__img" src="/static/img/icon-https.svg" alt="" aria-hidden="true" /> <div class="usa-media-block__body"> <p> <strong>Secure .gov websites use HTTPS</strong> <br /> A <strong>lock</strong> ( <span class="icon-lock"> <svg xmlns="http://www.w3.org/2000/svg" width="52" height="64" viewBox="0 0 52 64" class="usa-banner__lock-image" role="graphics-symbol" aria-labelledby="banner-lock-description" focusable="false"> <title id="banner-lock-title">Lock</title> <desc id="banner-lock-description"> Locked padlock icon </desc> <path fill="#000000" fill-rule="evenodd" d="M26 0c10.493 0 19 8.507 19 19v9h3a4 4 0 0 1 4 4v28a4 4 0 0 1-4 4H4a4 4 0 0 1-4-4V32a4 4 0 0 1 4-4h3v-9C7 8.507 15.507 0 26 0zm0 8c-5.979 0-10.843 4.77-10.996 10.712L15 19v9h22v-9c0-6.075-4.925-11-11-11z" /> </svg> </span>) or <strong>https://</strong> means you've safely connected to the .gov website. Share sensitive information only on official, secure websites. </p> </div> </div> </div> </div> </div> </section> <div class="usa-overlay"> </div> <header class="usa-header usa-header--extended usa-header--wide" data-testid="header" data-header > <div class="ncbi-header"> <div class="ncbi-header__container"> <a class="ncbi-header__logo-container" href="/"> <img alt=" PMC home page " class="ncbi-header__logo-image" src="/static/img/ncbi-logos/nih-nlm-ncbi--white.svg" /> </a>  <button type="button" class="usa-menu-btn ncbi-header__hamburger-button " aria-label="Show menu" data-testid="navMenuButton" > <svg aria-hidden="true" class="ncbi-hamburger-icon" fill="none" focusable="false" height="21" viewBox="0 0 31 21" width="31" xmlns="http://www.w3.org/2000/svg"> <path clip-rule="evenodd" d="M0.125 20.75H30.875V17.3333H0.125V20.75ZM0.125 12.2083H30.875V8.79167H0.125V12.2083ZM0.125 0.25V3.66667H30.875V0.25H0.125Z" fill="#F1F1F1" fill-rule="evenodd" /> </svg> </button>  <div class="ncbi-header__desktop-buttons">  <button type="button" class="usa-button usa-button--unstyled ncbi-header__desktop-button " aria-expanded="false" aria-controls="search-field-desktop-navigation" aria-label="Show search overlay" data-testid="toggleSearchPanelButton" data-toggle-search-panel-button > <svg class="usa-icon " role="graphics-symbol" aria-hidden="true" > <use xlink:href="/static/img/sprite.svg#search" /> </svg> Search </button>  <div class="ncbi-header__login-dropdown"> <button type="button" class="usa-button usa-button--unstyled ncbi-header__desktop-button ncbi-header__login-dropdown-button " aria-expanded="false" aria-controls="login-dropdown-menu" aria-label="Show login menu" data-testid="toggleLoginMenuDropdown" data-desktop-login-button > <svg class="usa-icon " role="graphics-symbol" aria-hidden="true" > <use xlink:href="/static/img/sprite.svg#person" /> </svg> <span data-login-dropdown-text>Log in</span>  <svg class="usa-icon ncbi-header__login-dropdown-icon ncbi-header__login-dropdown-icon--expand-less ncbi-header__login-dropdown-icon--hidden" role="graphics-symbol" aria-hidden="true" data-login-dropdown-up-arrow> <use xlink:href="/static/img/sprite.svg#expand_less" /> </svg>  <svg class="usa-icon ncbi-header__login-dropdown-icon ncbi-header__login-dropdown-icon--expand-more ncbi-header__login-dropdown-icon--hidden" role="graphics-symbol" aria-hidden="true" data-login-dropdown-down-arrow> <use xlink:href="/static/img/sprite.svg#expand_more" /> </svg> </button>  <ul class="usa-nav__submenu ncbi-header__login-dropdown-menu" id="login-dropdown-menu" data-desktop-login-menu-dropdown hidden> <li class="usa-nav__submenu-item">  <a href="https://www.ncbi.nlm.nih.gov/myncbi/" class="usa-link " > Dashboard </a> </li> <li class="usa-nav__submenu-item">  <a href="https://www.ncbi.nlm.nih.gov/myncbi/collections/bibliography/" class="usa-link " > Publications </a> </li> <li class="usa-nav__submenu-item">  <a href="https://www.ncbi.nlm.nih.gov/account/settings/" class="usa-link " > Account settings </a> </li> <li class="usa-nav__submenu-item"> <button type="button" class="usa-button usa-button--outline ncbi-header__login-dropdown-logout-button " data-testid="desktopLogoutButton" data-desktop-logout-button > Log out </button> </li> </ul> </div> </div> </div> </div>  <div class="ncbi-search-panel ncbi--show-only-at-desktop" data-testid="searchPanel" data-header-search-panel hidden> <div class="ncbi-search-panel__container"> <form action="https://www.ncbi.nlm.nih.gov/search/all/" aria-describedby="search-field-desktop-navigation-help-text" autocomplete="off" class="usa-search usa-search--big ncbi-search-panel__form" data-testid="form" method="GET" role="search"> <label class="usa-sr-only" data-testid="label" for="search-field-desktop-navigation"> Search… </label> <input class="usa-input" data-testid="textInput" id="search-field-desktop-navigation" name="term" placeholder="Search NCBI" type="search" value="" /> <button type="submit" class="usa-button " data-testid="button" > <span class="usa-search__submit-text"> Search NCBI </span> </button> </form> </div> </div> <nav aria-label="Primary navigation" class="usa-nav"> <p class="usa-sr-only" id="primary-navigation-sr-only-title"> Primary site navigation </p>  <button type="button" class="usa-nav__close ncbi-nav__close-button " aria-label="Close navigation menu" data-testid="navCloseButton" > <img src="/static/img/usa-icons/close.svg" alt="Close" /> </button>  <form class="usa-search usa-search--small ncbi--hide-at-desktop margin-top-6" role="search"> <label class="usa-sr-only" for="search-field"> Search </label> <input class="usa-input" id="search-field-mobile-navigation" type="search" placeholder="Search NCBI" name="search" /> <button type="submit" class="usa-button " >  <img src="data:image/svg+xml;base64,PHN2ZyB4bWxucz0iaHR0cDovL3d3dy53My5vcmcvMjAwMC9zdmciIGhlaWdodD0iMjQiIHZpZXdCb3g9IjAgMCAyNCAyNCIgd2lkdGg9IjI0Ij48cGF0aCBkPSJNMCAwaDI0djI0SDB6IiBmaWxsPSJub25lIi8+PHBhdGggZmlsbD0iI2ZmZiIgZD0iTTE1LjUgMTRoLS43OWwtLjI4LS4yN0E2LjQ3MSA2LjQ3MSAwIDAgMCAxNiA5LjUgNi41IDYuNSAwIDEgMCA5LjUgMTZjMS42MSAwIDMuMDktLjU5IDQuMjMtMS41N2wuMjcuMjh2Ljc5bDUgNC45OUwyMC40OSAxOWwtNC45OS01em0tNiAwQzcuMDEgMTQgNSAxMS45OSA1IDkuNVM3LjAxIDUgOS41IDUgMTQgNy4wMSAxNCA5LjUgMTEuOTkgMTQgOS41IDE0eiIvPjwvc3ZnPg==" class="usa-search__submit-icon" alt="Search" /> </button> </form>   <div class="ncbi-nav__mobile-login-menu ncbi--hide-at-desktop" data-mobile-login-menu hidden> <p class="ncbi-nav__mobile-login-menu-status"> Logged in as: <strong class="ncbi-nav__mobile-login-menu-email" data-mobile-login-email-text></strong> </p> <ul class="usa-nav__primary usa-accordion"> <li class="usa-nav__primary-item"> <a href="https://www.ncbi.nlm.nih.gov/myncbi/" class="usa-link " > Dashboard </a> </li> <li class="usa-nav__primary-item"> <a href="https://www.ncbi.nlm.nih.gov/myncbi/collections/bibliography/" class="usa-link " > Publications </a> </li> <li class="usa-nav__primary-item"> <a href="https://www.ncbi.nlm.nih.gov/account/settings/" class="usa-link " > Account settings </a> </li> </ul> </div> <button type="button" class="usa-button ncbi-nav__mobile-login-button ncbi--hide-at-desktop " data-testid="mobileLoginButton" data-mobile-login-button > Log in </button> </nav> </header> <section class="pmc-header pmc-header--basic" aria-label="PMC Header with search box"> <div class="pmc-nav-container"> <div class="pmc-header__bar"> <div class="pmc-header__logo"> <a href="/" title="Home" aria-label="PMC Home"></a> </div> <button type="button" class="usa-button usa-button--unstyled pmc-header__search__button" aria-label="Open search" data-ga-category="search" data-ga-action="PMC" data-ga-label="pmc_search_panel_mobile" > <svg class="usa-icon width-4 height-4 pmc-icon__open" aria-hidden="true" focusable="false" role="img"> <use xlink:href="/static/img/sprite.svg#search"></use> </svg> <svg class="usa-icon width-4 height-4 pmc-icon__close" aria-hidden="true" focusable="false" role="img"> <use xlink:href="/static/img/sprite.svg#close"></use> </svg> </button> </div> <div class="pmc-header__search"> <form class="usa-search usa-search--extra usa-search--article-right-column pmc-header__search__form" autocomplete="off" role="search"> <label class="usa-sr-only" for="pmc-search">Search PMC Full-Text Archive</label> <span class="autoComplete_wrapper flex-1"> <input class="usa-input width-full maxw-none" required="required" placeholder="Search PMC Full-Text Archive" id="pmc-search" type="search" name="term" data-autocomplete-url="/search/autocomplete/"/> </span> <button class="usa-button" type="submit" formaction="https://www.ncbi.nlm.nih.gov/pmc/" data-ga-category="search" data-ga-action="PMC" data-ga-label="PMC_search_button" > <span class="usa-search__submit-text">Search in PMC</span> <img src="/static/img/usa-icons-bg/search--white.svg" class="usa-search__submit-icon" alt="Search" /> </button> </form> <ul class="pmc-header__search__menu"> <li> <a class="usa-link" href="https://www.ncbi.nlm.nih.gov/pmc/advanced/" data-ga-action="featured_link" data-ga-label="advanced_search"> Advanced Search </a> </li> <li> <a class="usa-link" href="/journals/" data-ga-action="featured_link" data-ga-label="journal list"> Journal List </a> </li> <li> <a class="usa-link" href="/about/userguide/" data-ga-action="featured_link" data-ga-label="user guide"> User Guide </a> </li> </ul> </div> </div> </section> <div class="usa-section padding-top-0 desktop:padding-top-6 pmc-article-section" data-article-db="pmc" data-article-id="11213802"> <div class="grid-container pmc-actions-bar" aria-label="Actions bar" role="complementary"> <div class="grid-row"> <div class="grid-col-fill display-flex"> <div class="display-flex"> <ul class="usa-list usa-list--unstyled usa-list--horizontal"> <li class="margin-right-2 mobile-lg:margin-right-4 display-flex mob"> <button type="button" class="usa-button pmc-sidenav__container__open usa-button--unstyled width-auto display-flex" aria-label="Open resources" data-extra-class="is-visible-resources" data-ga-category="resources_accordion" data-ga-action="click" data-ga-label="mobile_icon" > <svg class="usa-icon width-4 height-4" aria-hidden="true" focusable="false" role="img"> <use xlink:href="/static/img/sprite.svg#more_vert"></use> </svg> </button> </li> <li class="margin-right-2 mobile-lg:margin-right-4 display-flex mob"> <a href="https://doi.org/10.1007/s10530-024-03291-9" class="usa-link display-flex" role="button" target="_blank" rel="noreferrer noopener" aria-label="View on publisher site" data-ga-category="actions" data-ga-action="click" data-ga-label="publisher_link_mobile" > <svg class="usa-icon width-4 height-4" aria-hidden="true" focusable="false" role="img"> <use xlink:href="/static/img/sprite.svg#launch"></use> </svg> </a> </li> <li class="margin-right-2 mobile-lg:margin-right-4 display-flex"> <a href="pdf/10530_2024_Article_3291.pdf" class="usa-link display-flex" role="button" aria-label="Download PDF" data-ga-category="actions" data-ga-action="click" data-ga-label="pdf_download_mobile" > <svg class="usa-icon width-4 height-4" aria-hidden="true" focusable="false" role="img"> <use xlink:href="/static/img/sprite.svg#file_download"></use> </svg> </a> </li> <li class="margin-right-2 mobile-lg:margin-right-4 display-flex"> <button class="usa-button usa-button--unstyled collections-dialog-trigger collections-button display-flex collections-button-empty" aria-label="Save article in MyNCBI collections." data-ga-category="actions" data-ga-action="click" data-ga-label="collections_button_mobile" data-collections-open-dialog-enabled="false" data-collections-open-dialog-url="https://account.ncbi.nlm.nih.gov/?back_url=https%3A%2F%2Fpmc.ncbi.nlm.nih.gov%2Farticles%2FPMC11213802%2F%23open-collections-dialog" data-in-collections="false" > <svg class="usa-icon width-4 height-4 usa-icon--bookmark-full" aria-hidden="true" focusable="false" role="img" hidden> <use xlink:href="/static/img/action-bookmark-full.svg#icon"></use> </svg> <svg class="usa-icon width-4 height-4 usa-icon--bookmark-empty" aria-hidden="true" focusable="false" role="img" hidden> <use xlink:href="/static/img/action-bookmark-empty.svg#icon"></use> </svg> </button> </li> <li class="margin-right-2 mobile-lg:margin-right-4 display-flex"> <button role="button" class="usa-button usa-button--unstyled citation-dialog-trigger display-flex" aria-label="Open dialog with citation text in different styles" data-ga-category="actions" data-ga-action="open" data-ga-label="cite_mobile" data-all-citations-url="/resources/citations/11213802/" data-citation-style="nlm" data-download-format-link="/resources/citations/11213802/export/" > <svg class="usa-icon width-4 height-4 usa-icon--bookmark-empty" aria-hidden="true" focusable="false" role="img" hidden> <use xlink:href="/static/img/sprite.svg#format_quote"></use> </svg> </button> </li> <li class="pmc-permalink display-flex"> <button type="button" class="usa-button usa-button--unstyled display-flex" aria-label="Show article permalink" aria-expanded="false" aria-haspopup="true" data-ga-category="actions" data-ga-action="open" data-ga-label="permalink_mobile" > <svg class="usa-icon width-4 height-4" aria-hidden="true" focusable="false" role="img"> <use xlink:href="/static/img/sprite.svg#share"></use> </svg> </button> <div class="pmc-permalink__dropdown" hidden> <div class="pmc-permalink__dropdown__container"> <h2 class="usa-modal__heading margin-top-0 margin-bottom-2">PERMALINK</h2> <div class="pmc-permalink__dropdown__content"> <input type="text" class="usa-input" value="https://pmc.ncbi.nlm.nih.gov/articles/PMC11213802/" aria-label="Article permalink"> <button class="usa-button display-inline-flex pmc-permalink__dropdown__copy__btn margin-right-0" title="Copy article permalink" data-ga-category="save_share" data-ga-action="link" data-ga-label="copy_link"> <svg class="usa-icon" aria-hidden="true" focusable="false" role="img"> <use xlink:href="/static/img/sprite.svg#content_copy"></use> </svg> <span class="margin-left-1">Copy</span> </button> </div> </div> </div> </li> </ul> </div> <button type="button" class="usa-button pmc-sidenav__container__open usa-button--unstyled width-auto display-flex" aria-label="Open article navigation" data-extra-class="is-visible-in-page" data-ga-category="actions" data-ga-action="open" data-ga-label="article_nav_mobile" > <svg class="usa-icon width-4 height-4" aria-hidden="true" focusable="false" role="img"> <use xlink:href="/static/img/sprite.svg#list"></use> </svg> </button> </div> </div> </div> <div class="grid-container desktop:padding-left-6"> <div id="article-container" class="grid-row grid-gap"> <div class="grid-col-12 desktop:grid-col-8 order-2 pmc-layout__content"> <div class="grid-container padding-left-0 padding-right-0"> <div class="grid-row desktop:margin-left-neg-6"> <div class="grid-col-12"> <div class="pmc-layout__disclaimer" role="complementary" aria-label="Disclaimer note"> As a library, NLM provides access to scientific literature. Inclusion in an NLM database does not imply endorsement of, or agreement with, the contents by NLM or the National Institutes of Health.<br/> Learn more: <a class="usa-link" data-ga-category="Link click" data-ga-action="Disclaimer" data-ga-label="New disclaimer box" href="/about/disclaimer/">PMC Disclaimer</a> | <a class="usa-link" data-ga-category="Link click" data-ga-action="PMC Copyright Notice" data-ga-label="New disclaimer box" href="/about/copyright/"> PMC Copyright Notice </a> </div> </div> </div> <div class="grid-row pmc-wm desktop:margin-left-neg-6">  <main id="main-content" class="usa-layout-docs__main usa-layout-docs grid-col-12 pmc-layout pmc-prose padding-0" > <section class="pmc-journal-banner text-center line-height-none" aria-label="Journal banner"><img src="https://cdn.ncbi.nlm.nih.gov/pmc/banners/logo-springeropen.png" alt="Springer logo" usemap="#pmc-banner-imagemap" width="500" height="75"><map name="pmc-banner-imagemap"><area alt="Link to Springer" title="Link to Springer" shape="default" href="https://doi.org/10.1007/s10530-024-03291-9" target="_blank" rel="noopener noreferrer"></map></section><article lang="en"><section aria-label="Article citation and metadata"><section class="pmc-layout__citation font-secondary font-xs"><div> <div class="display-inline-block"><button type="button" class="cursor-pointer text-no-underline bg-transparent border-0 padding-0 text-left margin-0 text-normal text-primary" aria-controls="journal_context_menu">Biol Invasions</button></div>. 2024 Apr 23;26(7):2037–2047. doi: <a href="https://doi.org/10.1007/s10530-024-03291-9" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">10.1007/s10530-024-03291-9</a> </div> <nav id="journal_context_menu" hidden="hidden"><ul class="menu-list font-family-ui" role="menu"> <li role="presentation"><a href="https://www.ncbi.nlm.nih.gov/pmc/?term=%22Biol%20Invasions%22%5Bjour%5D" class="usa-link" role="menuitem">Search in PMC</a></li> <li role="presentation"><a href="https://pubmed.ncbi.nlm.nih.gov/?term=%22Biol%20Invasions%22%5Bjour%5D" lang="en" class="usa-link" role="menuitem">Search in PubMed</a></li> <li role="presentation"><a href="https://www.ncbi.nlm.nih.gov/nlmcatalog?term=%22Biol%20Invasions%22%5BTitle%20Abbreviation%5D" class="usa-link" role="menuitem">View in NLM Catalog</a></li> <li role="presentation"><a href="?term=%22Biol%20Invasions%22%5Bjour%5D" class="usa-link" role="menuitem" data-add-to-search="true">Add to search</a></li> </ul></nav></section><section class="front-matter"><div class="ameta p font-secondary font-xs"> <hgroup><h1>Spatially limited pathogen pollution in an invasive tick and host system</h1></hgroup><div class="cg p"> <a href="https://pubmed.ncbi.nlm.nih.gov/?term=%22De%20Jesus%20CE%22%5BAuthor%5D" class="usa-link" aria-describedby="id1"><span class="name western">Carrie E De Jesus</span></a><div hidden="hidden" id="id1"> <h3><span class="name western">Carrie E De Jesus</span></h3> <div class="p"> <sup>1</sup>Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, FL USA </div> <div class="p">Find articles by <a href="https://pubmed.ncbi.nlm.nih.gov/?term=%22De%20Jesus%20CE%22%5BAuthor%5D" class="usa-link"><span class="name western">Carrie E De Jesus</span></a> </div> </div> <sup>1</sup>, <a href="https://pubmed.ncbi.nlm.nih.gov/?term=%22Harman%20MEA%22%5BAuthor%5D" class="usa-link" aria-describedby="id2"><span class="name western">Madison E A Harman</span></a><div hidden="hidden" id="id2"> <h3><span class="name western">Madison E A Harman</span></h3> <div class="p"> <sup>1</sup>Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, FL USA </div> <div class="p">Find articles by <a href="https://pubmed.ncbi.nlm.nih.gov/?term=%22Harman%20MEA%22%5BAuthor%5D" class="usa-link"><span class="name western">Madison E A Harman</span></a> </div> </div> <sup>1</sup>, <a href="https://pubmed.ncbi.nlm.nih.gov/?term=%22Sutton%20A%22%5BAuthor%5D" class="usa-link" aria-describedby="id3"><span class="name western">Amber Sutton</span></a><div hidden="hidden" id="id3"> <h3><span class="name western">Amber Sutton</span></h3> <div class="p"> <sup>1</sup>Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, FL USA </div> <div class="p">Find articles by <a href="https://pubmed.ncbi.nlm.nih.gov/?term=%22Sutton%20A%22%5BAuthor%5D" class="usa-link"><span class="name western">Amber Sutton</span></a> </div> </div> <sup>1</sup>, <a href="https://pubmed.ncbi.nlm.nih.gov/?term=%22Bredin%20S%22%5BAuthor%5D" class="usa-link" aria-describedby="id4"><span class="name western">Stephen Bredin</span></a><div hidden="hidden" id="id4"> <h3><span class="name western">Stephen Bredin</span></h3> <div class="p"> <sup>2</sup>Biology Department, George Mason University, Fairfax, VA USA </div> <div class="p">Find articles by <a href="https://pubmed.ncbi.nlm.nih.gov/?term=%22Bredin%20S%22%5BAuthor%5D" class="usa-link"><span class="name western">Stephen Bredin</span></a> </div> </div> <sup>2</sup>, <a href="https://pubmed.ncbi.nlm.nih.gov/?term=%22Romagosa%20CM%22%5BAuthor%5D" class="usa-link" aria-describedby="id5"><span class="name western">Christina M Romagosa</span></a><div hidden="hidden" id="id5"> <h3><span class="name western">Christina M Romagosa</span></h3> <div class="p"> <sup>1</sup>Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, FL USA </div> <div class="p">Find articles by <a href="https://pubmed.ncbi.nlm.nih.gov/?term=%22Romagosa%20CM%22%5BAuthor%5D" class="usa-link"><span class="name western">Christina M Romagosa</span></a> </div> </div> <sup>1</sup>, <a href="https://pubmed.ncbi.nlm.nih.gov/?term=%22Wisely%20SM%22%5BAuthor%5D" class="usa-link" aria-describedby="id6"><span class="name western">Samantha M Wisely</span></a><div hidden="hidden" id="id6"> <h3><span class="name western">Samantha M Wisely</span></h3> <div class="p"> <sup>1</sup>Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, FL USA </div> <div class="p">Find articles by <a href="https://pubmed.ncbi.nlm.nih.gov/?term=%22Wisely%20SM%22%5BAuthor%5D" class="usa-link"><span class="name western">Samantha M Wisely</span></a> </div> </div> <sup>1,</sup><sup>✉</sup> </div> <ul class="d-buttons inline-list"> <li><button class="d-button" aria-controls="aip_a" aria-expanded="false">Author information</button></li> <li><button class="d-button" aria-controls="anp_a" aria-expanded="false">Article notes</button></li> <li><button class="d-button" aria-controls="clp_a" aria-expanded="false">Copyright and License information</button></li> </ul> <div class="d-panels font-secondary-light"> <div id="aip_a" class="d-panel p" style="display: none"> <div class="p" id="Aff1"> <sup>1</sup>Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, FL USA </div> <div id="Aff2"> <sup>2</sup>Biology Department, George Mason University, Fairfax, VA USA </div> <div class="author-notes p"><div class="fn" id="_fncrsp93pmc__"> <sup>✉</sup><p class="display-inline">Corresponding author.</p> </div></div> </div> <div id="anp_a" class="d-panel p" style="display: none"><div class="notes p"><section id="historyarticle-meta1" class="history"><p>Received 2023 Feb 20; Accepted 2024 Mar 7; Issue date 2024.</p></section></div></div> <div id="clp_a" class="d-panel p" style="display: none"> <div>© The Author(s) 2024</div> <p><strong>Open Access</strong> This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit <a href="https://creativecommons.org/licenses/by/4.0/" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">http://creativecommons.org/licenses/by/4.0/</a>.</p> <div class="p"><a href="/about/copyright/" class="usa-link">PMC Copyright notice</a></div> </div> </div> <div>PMCID: PMC11213802 PMID: <a href="https://pubmed.ncbi.nlm.nih.gov/38947421/" class="usa-link">38947421</a> </div> </div></section></section><section aria-label="Article content"><section class="body main-article-body"><section class="abstract" id="Abs1"><h2>Abstract</h2> <p id="Par1">Expansion of global commerce has facilitated pathogen pollution via the transportation and translocation of invasive species and their associated parasites and pathogens. In Florida, imported cane toads (<em>Rhinella horribilis</em>) were accidentally and intentionally released on multiple occasions. Early populations were found to be infested with the invasive tick, <em>Amblyomma rotundatum</em>, yet it is unknown if these ticks dispersed with their hosts as cane toads spread throughout much of the state. The objectives of our investigation were to (1) determine if there are fewer tick infestations on toads at the periphery than at the core of their distribution as predicted by founder effect events, and (2) identify if ticks were infected with exotic pathogens. We captured toads from 10 populations across Florida. We collected ticks, vent tissue, and tick attachment site tissue from each toad, then tested samples for bacteria in the genus, <em>Rickettsia</em>. We found that 3/10 populations had toads that were infested with <em>A. rotundatum</em>, and infested individuals were in the earliest introduced populations at the core of their distribution. Pathogen testing confirmed <em>Rickettisa bellii</em> in ticks, but not in toad tissues. Haplotype networks could not clearly distinguish if <em>R. bellii</em> in Florida was more closely related to North or South American strains, but host-tick associations suggest that the pathogen was exotic to Florida. Our investigation demonstrated that an invasive species facilitated the introduction of parasites and pathogens into Florida, yet the invasive tick species encountered limitations to dispersal on this host species.</p> <section id="sec1"><h3 class="pmc_sec_title">Supplementary Information</h3> <p>The online version contains supplementary material available at 10.1007/s10530-024-03291-9.</p></section><section id="kwd-group1" lang="en" class="kwd-group"><p><strong>Keywords:</strong> <em>Amblyomma rotundatum</em>, Cane toad, Core-periphery distribution, Enemy release hypothesis, Founder effects, Invasive species, <em>Rickettsia bellii</em>, <em>Rhinella marina</em>, <em>Rhinella horribilis</em></p></section></section><section id="Sec1"><h2 class="pmc_sec_title">Introduction</h2> <p id="Par2">The expansion of global trade and travel has created pathways for invasive species and accelerated their spread worldwide (Sakai et al. <a href="#CR51" class="usa-link" aria-describedby="CR51">2001</a>; Van Kleunen et al. <a href="#CR55" class="usa-link" aria-describedby="CR55">2015</a>; Capinha et al. <a href="#CR7" class="usa-link" aria-describedby="CR7">2017</a>). Once introduced, species with traits that support rapid reproduction and dispersal can become successful invaders. Whether intentional or accidental, these introductions can directly or indirectly alter invaded ecosystems resulting in decreased biodiversity, ecosystem function, and ecosystem services (Floerl et al. <a href="#CR15" class="usa-link" aria-describedby="CR15">2009</a>; Mooney and Cleland <a href="#CR39" class="usa-link" aria-describedby="CR39">2001</a>).</p> <p id="Par3">One way invasive species alter ecosystems is via the co-introduction of non-native parasites and pathogens, i.e. pathogen pollution (Cunningham et al. <a href="#CR10" class="usa-link" aria-describedby="CR10">2003</a>). An emerging focus of invasion ecology is the study of how host-parasite dynamics change with the introduction of invasive species to a novel area. Parasites and pathogens are ubiquitous in biological communities and can influence both the invasion success of non-native species and the fitness of native species (Prenter et al. <a href="#CR48" class="usa-link" aria-describedby="CR48">2004</a>). Parasites and pathogens can negatively impact the fecundity of hosts via energy trade-offs that allocate the host’s energy to immune response instead of dispersal, growth, or reproduction (Luong et al. <a href="#CR34" class="usa-link" aria-describedby="CR34">2017</a>), in turn regulating host population levels (Prenter et al. <a href="#CR48" class="usa-link" aria-describedby="CR48">2004</a>).</p> <p id="Par4">Following the establishment of an invasive host species, parasites and pathogens may be distributed unevenly across the invasive host range due to a variety of ecological processes. Peripheral host populations, those that are on the leading edge of the expansion front, may have lower rates of parasitism due to the stochastic infestation rates of immigrant individuals which may lead to lower rates of parasitism in colonizing populations (Phillips et al. <a href="#CR45" class="usa-link" aria-describedby="CR45">2010</a>; Barnett et al. <a href="#CR3" class="usa-link" aria-describedby="CR3">2018</a>). Alternatively, but not mutually exclusively, core host populations, those that established early in the invasion process, may have more parasites than populations along the expanding peripheral edge because habitat at the site of establishment may be of higher quality than habitat in isolated, peripheral populations, leading to higher host and parasite densities at the distribution core. In addition, cross-species transmission of parasites from native or previously established non-native species to the invading species (spill-back) may also facilitate uneven or patchy distribution of parasitism across the landscape (Chalkowski et al. <a href="#CR8" class="usa-link" aria-describedby="CR8">2018</a>). Thus, patterns of parasite distribution on invasive host species can provide insight into the underlying processes driving host-parasite relationships.</p> <p id="Par5">While some parasite populations may struggle to persist during the invasion process, those that successfully become established often share generalist phenotypic traits. Parasites that are host generalists (Ewen et al. <a href="#CR14" class="usa-link" aria-describedby="CR14">2012</a>) can tolerate variable climatic conditions (Polo et al. <a href="#CR47" class="usa-link" aria-describedby="CR47">2021</a>) and typically have simple life cycles that require few host species or can reproduce asexually. Many tick species (<em>Ixodida</em>) exemplify these traits and are frequently successful invaders (Barré and Uilenberg <a href="#CR4" class="usa-link" aria-describedby="CR4">2010</a>). Multiple tick species have successfully invaded and established in the United States (Burridge <a href="#CR5" class="usa-link" aria-describedby="CR5">2011</a>) including <em>Amblyomma rotundatum,</em> which originated from Central and South America. <em>Amblyomma rotundatum</em> is a 3-host tick species, yet the same host species may support all three tick life stages. <em>A. rotundatum</em> reproduces parthenogenically, which streamlines population growth by eliminating the need to search for a mate. <em>Amblyomma rotundatum</em> is the one of the most prolific generalist feeders reported in the literature, with 59 reported herpetological host species (De Jesus <a href="#CR20" class="usa-link" aria-describedby="CR20">2021</a>) and 16 mammal host species (Guglielmone and Nava <a href="#CR16" class="usa-link" aria-describedby="CR16">2010</a>). This combination of traits has made it a highly successful invasive species.</p> <p id="Par6">While it has been hypothesized that <em>A. rotundatum</em> was introduced into Florida with its native host, the cane toad (<em>Rhinella horribilis</em> formerly <em>R. marina</em>, Acevedo et al. <a href="#CR1" class="usa-link" aria-describedby="CR1">2016</a>, Mittan-Moreau et al. <a href="#CR38" class="usa-link" aria-describedby="CR38">2022</a>, Oliver et al. <a href="#CR42" class="usa-link" aria-describedby="CR42">1993</a>), the origins of <em>A. rotundatum</em> in Florida are not well understood and are likely complicated. Multiple invasive reptile species that originated from Central and South America have been introduced in the decades after cane toads became established (Krysko et al. <a href="#CR25" class="usa-link" aria-describedby="CR25">2011</a>; Nava et al. <a href="#CR40" class="usa-link" aria-describedby="CR40">2017</a>) providing additional potential source populations of <em>A. rotundatum</em>, and other non-native reptiles including the Burmese python (<em>Python bivittatus</em>) from Southeast Asia and the Peter’s Rock agama (<em>Agama picticauda</em>) from Africa (Corn et al. <a href="#CR9" class="usa-link" aria-describedby="CR9">2014</a>, pers. obs.) have been observed in Florida with <em>A. rotundatum</em> infestations<em>.</em> Additionally, cases of parasite spillover of <em>A. rotundatum</em> have been reported on native snake species like the southern black racer (<em>Coluber constrictor priapus</em>) in southernmost Florida and cottonmouth (<em>Agkistrodon piscivorus conanti</em>) (Hanson et al. <a href="#CR18" class="usa-link" aria-describedby="CR18">2007</a>; Corn et al. <a href="#CR9" class="usa-link" aria-describedby="CR9">2014</a>; Lillywhite and Sheehy <a href="#CR33" class="usa-link" aria-describedby="CR33">2019</a>) from northwest peninsular Florida suggesting that this invasive tick species is distributed throughout Florida.</p> <p id="Par7">Cane toads are an infamous invasive species that were introduced into subtropical and tropical climates worldwide in failed biological control attempts for sugar cane beetles (King and Krakauer <a href="#CR22" class="usa-link" aria-describedby="CR22">1966</a>; Lever <a href="#CR31" class="usa-link" aria-describedby="CR31">2001</a>). Cane toads were originally introduced into Florida cane sugar fields in Palm Beach County during the 1930s and 1940s, although those populations did not become established (King and Krakauer <a href="#CR22" class="usa-link" aria-describedby="CR22">1966</a>). Instead, cane toads likely became established in Florida in the 1950s after an accidental mass-release of toads imported to the Miami Airport from Colombia (Krakauer <a href="#CR23" class="usa-link" aria-describedby="CR23">1968</a>; Oliver et al. <a href="#CR42" class="usa-link" aria-describedby="CR42">1993</a>). Then in the 1960s, animal dealers intentional released additional cane toads from Colombia in Pembroke Park near Ft. Lauderdale in Broward County and from Suriname in Kendall near Miami in Miami-Dade County (King and Krakauer <a href="#CR22" class="usa-link" aria-describedby="CR22">1966</a>). Thus, two populations were established approximately 60 km from each other in southeast Florida (Fig. <a href="#Fig1" class="usa-link">1</a>). Since these introductions, cane toads have utilized canals and urbanized habitats to establish expanding populations across southern Florida and northwards into central Florida (Meshaka et al. <a href="#CR37" class="usa-link" aria-describedby="CR37">2006</a>; Wilson <a href="#CR57" class="usa-link" aria-describedby="CR57">2016</a>). <em>Amblyomma rotundatum</em> were first identified infesting invasive cane toads in Miami, Florida in the late 1970’s (Oliver et al. <a href="#CR42" class="usa-link" aria-describedby="CR42">1993</a>).</p> <figure class="fig xbox font-sm" id="Fig1"><h3 class="obj_head">Fig. 1.</h3> <p class="img-box line-height-none margin-x-neg-2 tablet:margin-x-0 text-center"><a class="tileshop" target="_blank" href="https://www.ncbi.nlm.nih.gov/core/lw/2.0/html/tileshop_pmc/tileshop_pmc_inline.html?title=Click%20on%20image%20to%20zoom&p=PMC3&id=11213802_10530_2024_3291_Fig1_HTML.jpg"><img class="graphic zoom-in" src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/b14e/11213802/cb1a9e6bb721/10530_2024_3291_Fig1_HTML.jpg" loading="lazy" id="MO1" height="443" width="696" alt="Fig. 1"></a></p> <div class="p text-right font-secondary"><a href="figure/Fig1/" class="usa-link" target="_blank" rel="noopener noreferrer">Open in a new tab</a></div> <figcaption><p>Circles indicate field sites where toads were collected. Blue circles identify sampling sites near introduction locations in Florida. Parentheses display number of individual toads infected with ticks</p></figcaption></figure><p id="Par8">Ticks are associated with numerous bacterial pathogens. Bacteria in the genus <em>Rickettsia</em> are the most ubiquitous tick-borne pathogen reported globally (Parola et al. <a href="#CR43" class="usa-link" aria-describedby="CR43">2013</a>; Piotrowski and Rymaszewska <a href="#CR46" class="usa-link" aria-describedby="CR46">2020</a>). <em>Rickettsia</em> bacteria vary in their pathogenicity from causing severe human and animal illnesses to nonpathogenic endosymbionts (Parola et al. <a href="#CR43" class="usa-link" aria-describedby="CR43">2013</a>; El Karkouri et al. <a href="#CR13" class="usa-link" aria-describedby="CR13">2022</a>). <em>Rickettsia</em> are likely candidates for pathogen pollution because of their unique transmission properties. Unlike some tick-borne pathogens, <em>Rickettsia</em> can be transmitted transovarially, i.e., adult female ticks transmit <em>Rickettsia</em> bacteria to their eggs and the resulting larvae emerge infected (Horta et al. <a href="#CR19" class="usa-link" aria-describedby="CR19">2006</a>; Laukaitis and Macaluso <a href="#CR29" class="usa-link" aria-describedby="CR29">2021</a>). The subsequent life stages can then maintain that infection between molts, referred to as transstadial transmission (Labruna <a href="#CR27" class="usa-link" aria-describedby="CR27">2009</a>; Parola et al. <a href="#CR43" class="usa-link" aria-describedby="CR43">2013</a>). Transovarial and transstadial bacterial transmission facilitate transmission from one life stage to another without the tick taking a bloodmeal from an infected host; therefore pathogens can be imported in infected ticks even if those ticks are infesting an uninfected host. All life stages disseminate <em>Rickettsia</em> (Parola et al. <a href="#CR43" class="usa-link" aria-describedby="CR43">2013</a>), although some species of <em>Rickettsia</em> require infected hosts to maintain infection in the tick population. Nonetheless, transovarial transmission provides exotic rickettsial species with fewer barriers to overcome during the invasion process than bacteria with other transmission routes.</p> <p id="Par9"><em>Rickettsia</em> species have been reported in <em>A</em>. <em>rotundatum</em> within the cane toad’s native range. One commonly reported species is <em>Rickettsia bellii</em> (Labruna et al. <a href="#CR28" class="usa-link" aria-describedby="CR28">2004</a>; Luz et al. <a href="#CR36" class="usa-link" aria-describedby="CR36">2018</a>; Sánchez-Montes et al. <a href="#CR52" class="usa-link" aria-describedby="CR52">2019</a>) which has been reported in 19 tick species across the Americas and is considered nonpathogenic (Krawczak et al. <a href="#CR24" class="usa-link" aria-describedby="CR24">2018</a>). <em>Rickettsia bellii</em> infections are maintained by both transovarial and transstadial transmission. This species also has distinct clades for North and South American strains (Krawczak et al. <a href="#CR24" class="usa-link" aria-describedby="CR24">2018</a>). <em>Rickettsia bellii</em> provides a unique model system to observe how <em>Rickettsia</em> respond to the biological invasions processes of their tick host.</p> <p id="Par10">Cane toads; the tick, <em>Amblyomma rotundatum</em>; and the bacterial species, <em>Rickettsia belli</em> provide a unique host-parasite-microbe system to empirically observe geographic patterns of parasite infestation and microbe infection that inform parasite ecology within the distribution of an invasive species. The objectives of our investigation were to (1) determine if core-periphery patterns of tick infestation rates occurred in cane toads. We predicted that there would be fewer tick infestations in toads at the periphery of the invasion than at the core of their distribution and (2) identify whether these invasive ticks were infected with exotic <em>Rickettsia</em> species not native to Florida and (3) determine if the pattern of a core-periphery distribution was maintained in the exotic bacteria found in ticks or cane toads. To conduct this study, we surveyed cane toad populations along a core to periphery distributional gradient for the presence of ticks. We then screened ticks and cane toads for infection with rickettsial bacteria.</p></section><section id="Sec2"><h2 class="pmc_sec_title">Materials and methods</h2> <section id="Sec3"><h3 class="pmc_sec_title">Cane toad and tick collection</h3> <p id="Par11">We collected cane toads from 10 populations across their invasive range in Florida (Fig. <a href="#Fig1" class="usa-link">1</a>) from April–June 2021. Sampling dates overlapped with the cane toad breeding season, which occurs from March through September in Florida (Krakauer <a href="#CR23" class="usa-link" aria-describedby="CR23">1968</a>). We determined survey locations based on prior studies and citizen science reports through EDDMapS (Mittan & Zamudio <a href="#CR100" class="usa-link" aria-describedby="CR100">2019</a>; Rubio et al. <a href="#CR200" class="usa-link" aria-describedby="CR200">2020</a>; EDDMapS <a href="#CR201" class="usa-link" aria-describedby="CR201">2024</a>). We conducted surveys near water bodies in urban areas at night, when and where cane toads actively forage (Wilson <a href="#CR57" class="usa-link" aria-describedby="CR57">2016</a>). We captured toads by hand and placed them into individual ventilated plastic containers until ticks could be removed; animals were held in containers for < 24 h (University of Florida IACUC #202111387).</p> <p id="Par12">Toads were visually inspected for all life stages of attached ticks prior to euthanasia. If ticks were present, they were removed from the toad using fine tip forceps. Once removed, ticks were stored in 100% molecular-grade ethanol until they could be identified and extracted for DNA. We identified ticks to species morphologically using taxonomic keys from the United States, Central America, and South America (Keirans and Oliver <a href="#CR21" class="usa-link" aria-describedby="CR21">1993</a>; Guzman-Cornejo et al. <a href="#CR17" class="usa-link" aria-describedby="CR17">2011</a>; Nava et al. <a href="#CR40" class="usa-link" aria-describedby="CR40">2017</a>).</p> <p id="Par13">After ticks were removed, toads were humanely euthanized through dermal application of 20% benzocaine gel (American Veterinary Medical Association <a href="#CR2" class="usa-link" aria-describedby="CR2">2020</a>, IACUC #202111387). Following euthanasia, we collected morphological data (i.e., snout-urostyle length (SUL), mass, and sex) and tissue biopsies from all toads. When ticks were found on the toads, we took a skin biopsy at the site of tick attachment, referred to hereafter as attachment tissue. We collected vent tissue from all toads to assess if <em>Rickettsia</em> bacteria could be collected from highly vascularized tissues (Levin et al. <a href="#CR32" class="usa-link" aria-describedby="CR32">2016</a>). Tissue samples were placed in 100% molecular grade ethanol and then stored at − 20 °C until they were extracted for DNA.</p></section><section id="Sec4"><h3 class="pmc_sec_title">DNA extractions</h3> <p id="Par14">Tick and tissue samples were rinsed once with PBS buffer and twice with DI water to remove debris or benzocaine gel before DNA was extracted. DNA was then extracted from ticks and cane toad tissue samples using the Qiagen Gentra Puregene Kit (Valenica, CA, USA) with the manufacturer protocol. We extracted DNA from individual adults and nymphal ticks, but aggregated larvae into pools of 25 ticks per host. We cut vent and attachment site tissue into 10 ng pieces before extraction (Qiagen <a href="#CR49" class="usa-link" aria-describedby="CR49">2014</a>). We stored eluted DNA at − 20 °C until PCR protocols were conducted.</p></section><section id="Sec5"><h3 class="pmc_sec_title">PCR and sequencing</h3> <p id="Par15">To screen for <em>Rickettsia,</em> we initially targeted the gltA gene, as it can broadly detect <em>Rickettsia</em> species (Roux et al. <a href="#CR50" class="usa-link" aria-describedby="CR50">1997</a>). We further analyzed <em>Rickettsia</em> positive samples using primers that amplified portions of two additional genes: atpA and coxA because they have previously been shown to successfully amplify <em>R. bellii</em> (Weinert et al. <a href="#CR56" class="usa-link" aria-describedby="CR56">2009</a>; Krawczak et al. <a href="#CR24" class="usa-link" aria-describedby="CR24">2018</a>). All three genes were used to differentiate between strains of <em>R. bellii</em>. We ran a positive and negative control for each PCR assay. We used a positive control from a <em>Rickettsia</em> sp. collected from an <em>Ixodes scapularis</em> specimen. Our negative control was PCR grade water. All PCR products were run on a 1.5% agarose gel with RedView Stain (Genecopoeia, Rockville, MD) and visualized on UVP gel documentation system (Analytik-Jena, Beverly, MA). We considered samples positive if they had the appropriate band size. All PCR products were cleaned with SAP/Exonuclease and sent to a commercial lab for Sanger sequencing (Functional Biosciences, WI, USA). Consensus sequences were constructed from forward and reverse primers for atpA and gltA. For coxA only forward sequences were used due to a bacterial coinfection of <em>Chryseobacterium sp</em>. that was co-amplified with the reverse primer (Supplementary Materials). Sequences were then aligned using Geneious software (2019.1.3) (Biomatters Ltd., Auckland, New Zealand) and compared to sequences in GenBank using NCBI BLAST.</p></section><section id="Sec6"><h3 class="pmc_sec_title">Phylogenetic and network analyses</h3> <p id="Par16">We concatenated and assembled our three gene sequences in Geneious and compared them with sequence data from <em>R. bellii</em> isolates and field collected specimens deposited in GenBank (<a href="https://www.ncbi.nlm.nih.gov/nuccore/?term=OP650115:OP650207%5Bpacc%5D" class="usa-link" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">OP650115–OP650207</a>). We aligned our samples with GenBank samples using ClustalW (Thompson et al. <a href="#CR53" class="usa-link" aria-describedby="CR53">1994</a>) in MEGA X (Kumar et al. <a href="#CR26" class="usa-link" aria-describedby="CR26">2018</a>) software. Each alignment was visually examined to make sure all sequences aligned.</p> <p id="Par17">To determine the relationship between our <em>Rickettsia</em> samples and other Rickettisial species, we constructed a tree using the Tamura-Nei genetic distance model and the UPGMA tree building method with 500 bootstrap. We then examined each target gene and all 3 genes concatenated in a minimum spanning network in PopArt (Leigh and Bryant <a href="#CR30" class="usa-link" aria-describedby="CR30">2015</a>) in order to compare our samples from haplotypes in North America and South America. Additional sequences from Krawczak et al. (<a href="#CR24" class="usa-link" aria-describedby="CR24">2018</a>) and GenBank (Supplementary Table <a href="#MOESM1" class="usa-link">1</a>) supplemented the haplotype network.</p></section></section><section id="Sec7"><h2 class="pmc_sec_title">Results</h2> <section id="Sec8"><h3 class="pmc_sec_title">Cane toad and tick collections</h3> <p id="Par18">We collected a total of 234 cane toads from 10 populations across south and central Florida. Surveys yielded 54 females and 180 males, including 212 adult toads and 22 juveniles. Snout-urostyle length ranged from 6.8 to 13.3 cm with an average of 10.8 ± 1.6 cm. Of all cane toads collected, 56/234 (23.9%) toads were infested with ticks (Table <a href="#Tab1" class="usa-link">1</a>). Toads that were infested with ticks were collected from 3/10 populations: Homestead, Miami, and West Palm Beach (Fig. <a href="#Fig1" class="usa-link">1</a>). Among the three infested toad populations, ticks were found on toads 60.0% (57/95 toads) of the time. All ticks were identified as <em>A. rotundatum</em>. In total we collected 495 <em>A. rotundatum</em> and toads were infested with 5.2 ± 1.5 ticks/toad (Table <a href="#Tab1" class="usa-link">1</a>).</p> <section class="tw xbox font-sm" id="Tab1"><h4 class="obj_head">Table 1.</h4> <div class="caption p"><p>Total number of ticks collected at each field site by life stage and average tick infestation per toad based on site. Tick infestation average is based only on cane toad populations with ticks. Standard error is reported for average tick infestation</p></div> <div class="tbl-box p" tabindex="0"><table class="content" frame="hsides" rules="groups"> <thead><tr> <th align="left" colspan="1" rowspan="1">Field site (N)</th> <th align="left" colspan="1" rowspan="1">Larvae</th> <th align="left" colspan="1" rowspan="1">Nymph</th> <th align="left" colspan="1" rowspan="1">Adult</th> <th align="left" colspan="1" rowspan="1">Total ticks collected</th> <th align="left" colspan="1" rowspan="1">Average tick infestation</th> </tr></thead> <tbody> <tr> <td align="left" colspan="1" rowspan="1">West Palm Beach (35)</td> <td align="left" colspan="1" rowspan="1">41</td> <td align="left" colspan="1" rowspan="1">24</td> <td align="left" colspan="1" rowspan="1">9</td> <td align="left" colspan="1" rowspan="1">74</td> <td align="left" colspan="1" rowspan="1">2.3 (± 0.7)</td> </tr> <tr> <td align="left" colspan="1" rowspan="1">Miami (21)</td> <td align="left" colspan="1" rowspan="1">16</td> <td align="left" colspan="1" rowspan="1">16</td> <td align="left" colspan="1" rowspan="1">8</td> <td align="left" colspan="1" rowspan="1">40</td> <td align="left" colspan="1" rowspan="1">1.9 (± 0.7)</td> </tr> <tr> <td align="left" colspan="1" rowspan="1">Homestead (39)</td> <td align="left" colspan="1" rowspan="1">307</td> <td align="left" colspan="1" rowspan="1">66</td> <td align="left" colspan="1" rowspan="1">8</td> <td align="left" colspan="1" rowspan="1">381</td> <td align="left" colspan="1" rowspan="1">10.2 (± 3.2)</td> </tr> <tr> <td align="left" colspan="1" rowspan="1">Total (75)</td> <td align="left" colspan="1" rowspan="1">364</td> <td align="left" colspan="1" rowspan="1">106</td> <td align="left" colspan="1" rowspan="1">25</td> <td align="left" colspan="1" rowspan="1">495</td> <td align="left" colspan="1" rowspan="1">5.2 (± 1.5)</td> </tr> </tbody> </table></div> <div class="p text-right font-secondary"><a href="table/Tab1/" class="usa-link" target="_blank" rel="noopener noreferrer">Open in a new tab</a></div></section></section><section id="Sec9"><h3 class="pmc_sec_title">Rickettsia screening</h3> <p id="Par19">We tested ticks and tissue from vent and attachment sites for the presence of <em>Rickettsia</em> bacteria. Of the toads infested with ticks, 27/56 toads (48.2%) had ticks infected with <em>Rickettsia</em> (Table <a href="#Tab2" class="usa-link">2</a>). Homestead, Miami, and West Palm Beach all had ticks with rickettsial infections. We did not detect <em>Rickettsia</em> in any vent or tick attachment tissue samples.</p> <section class="tw xbox font-sm" id="Tab2"><h4 class="obj_head">Table 2.</h4> <div class="caption p"><p>Toads with ticks infected with <em>Rickettsia</em> bacteria based on gltA gene. Wilson Score 95% confidence interval was calculated for pathogen prevalence of <em>Rickettsia</em></p></div> <div class="tbl-box p" tabindex="0"><table class="content" frame="hsides" rules="groups"> <thead><tr> <th align="left" colspan="1" rowspan="1">Field sites with ticks</th> <th align="left" colspan="1" rowspan="1">Toads with <em>Rickettsia</em> positive ticks</th> </tr></thead> <tbody> <tr> <td align="left" colspan="1" rowspan="1">West Palm Beach</td> <td align="left" colspan="1" rowspan="1">7/15 (47%) (± 0.23)</td> </tr> <tr> <td align="left" colspan="1" rowspan="1">Miami</td> <td align="left" colspan="1" rowspan="1">1/9 (11%) (± 0.21)</td> </tr> <tr> <td align="left" colspan="1" rowspan="1">Homestead</td> <td align="left" colspan="1" rowspan="1">19/32 (59%) (± 0.16)</td> </tr> <tr> <td align="left" colspan="1" rowspan="1">Total</td> <td align="left" colspan="1" rowspan="1">27/56 (48%) (± 0.05)</td> </tr> </tbody> </table></div> <div class="p text-right font-secondary"><a href="table/Tab2/" class="usa-link" target="_blank" rel="noopener noreferrer">Open in a new tab</a></div></section></section><section id="Sec10"><h3 class="pmc_sec_title">Phylogenetic and haplotype network analyses</h3> <p id="Par20">Examining the gltA gene we found that our samples grouped with <em>R. bellii</em> and not with other rickettsial species (Fig. <a href="#Fig2" class="usa-link">2</a>). We found that all three target genes from <em>A. rotundatum</em> also closely matched with <em>R. bellii</em> sequences from both North and South America (Table <a href="#Tab3" class="usa-link">3</a>).</p> <figure class="fig xbox font-sm" id="Fig2"><h4 class="obj_head">Fig. 2.</h4> <p class="img-box line-height-none margin-x-neg-2 tablet:margin-x-0 text-center"><a class="tileshop" target="_blank" href="https://www.ncbi.nlm.nih.gov/core/lw/2.0/html/tileshop_pmc/tileshop_pmc_inline.html?title=Click%20on%20image%20to%20zoom&p=PMC3&id=11213802_10530_2024_3291_Fig2_HTML.jpg"><img class="graphic zoom-in" src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/b14e/11213802/e1b4a72562d5/10530_2024_3291_Fig2_HTML.jpg" loading="lazy" id="MO2" height="702" width="696" alt="Fig. 2"></a></p> <div class="p text-right font-secondary"><a href="figure/Fig2/" class="usa-link" target="_blank" rel="noopener noreferrer">Open in a new tab</a></div> <figcaption><p>A phylogenetic tree for the gltA gene of <em>Rickettsia</em> species was constructed using the Tamura-Nei genetic distance model and the UPGMA tree building method with 500 bootstrap. Sequences from this study are designated as Tick #. Previously published sequences are listed by <em>Rickettsia</em> species and accession number. Blue dots indicate R. bellii collected from South America, and red dots are North America</p></figcaption></figure><section class="tw xbox font-sm" id="Tab3"><h4 class="obj_head">Table 3.</h4> <div class="caption p"><p>Closest matches in GenBank to our <em>A. rotundatum</em> samples for Florida for each target gene</p></div> <div class="tbl-box p" tabindex="0"><table class="content" frame="hsides" rules="groups"> <thead><tr> <th align="left" colspan="1" rowspan="1">Pathogen species</th> <th align="left" colspan="1" rowspan="1">Target gene</th> <th align="left" colspan="1" rowspan="1">% Identical (Accession #)</th> <th align="left" colspan="1" rowspan="1">Tick species</th> <th align="left" colspan="1" rowspan="1">Loc locality</th> </tr></thead> <tbody> <tr> <td align="left" colspan="1" rowspan="1"><em>Rickettsia bellii</em></td> <td align="left" colspan="1" rowspan="1">atpA</td> <td align="left" colspan="1" rowspan="1">99.9%(<a href="https://www.ncbi.nlm.nih.gov/nuccore/MT009131" class="usa-link" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">MT009131</a>)</td> <td align="left" colspan="1" rowspan="1"><em>Amblyomma dissimile</em></td> <td align="left" colspan="1" rowspan="1">French Guiana</td> </tr> <tr> <td align="left" colspan="1" rowspan="1"><em>Rickettsia bellii</em></td> <td align="left" colspan="1" rowspan="1">coxA</td> <td align="left" colspan="1" rowspan="1">99.7%(<a href="https://www.ncbi.nlm.nih.gov/nuccore/CP000849" class="usa-link" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">CP000849</a>)</td> <td align="left" colspan="1" rowspan="1"><em>Dermacentor variabilis</em></td> <td align="left" colspan="1" rowspan="1">Ohio, USA</td> </tr> <tr> <td align="left" colspan="1" rowspan="1"><em>Rickettsia bellii</em></td> <td align="left" colspan="1" rowspan="1">gltA</td> <td align="left" colspan="1" rowspan="1">99.4% (<a href="https://www.ncbi.nlm.nih.gov/nuccore/MW384865" class="usa-link" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">MW384865</a>)</td> <td align="left" colspan="1" rowspan="1"><em>Amblyomma ovale</em></td> <td align="left" colspan="1" rowspan="1">El Salvador</td> </tr> </tbody> </table></div> <div class="p text-right font-secondary"><a href="table/Tab3/" class="usa-link" target="_blank" rel="noopener noreferrer">Open in a new tab</a></div></section><p id="Par21">Using the minimum spanning network based on the atpA gene, we found that our samples shared the same haplotype as two samples from South America detected in <em>Amblyomma dissmile</em> (Fig. <a href="#Fig3" class="usa-link">3</a>) (Supplementary Materials Table 1). However, there was not a clear distinction between North and South American haplotypes for any sequence or when all three sequences were concatenated (Fig. <a href="#Fig3" class="usa-link">3</a>).</p> <figure class="fig xbox font-sm" id="Fig3"><h4 class="obj_head">Fig. 3.</h4> <p class="img-box line-height-none margin-x-neg-2 tablet:margin-x-0 text-center"><a class="tileshop" target="_blank" href="https://www.ncbi.nlm.nih.gov/core/lw/2.0/html/tileshop_pmc/tileshop_pmc_inline.html?title=Click%20on%20image%20to%20zoom&p=PMC3&id=11213802_10530_2024_3291_Fig3_HTML.jpg"><img class="graphic zoom-in" src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/b14e/11213802/947e6d9d23dc/10530_2024_3291_Fig3_HTML.jpg" loading="lazy" id="MO3" height="612" width="696" alt="Fig. 3"></a></p> <div class="p text-right font-secondary"><a href="figure/Fig3/" class="usa-link" target="_blank" rel="noopener noreferrer">Open in a new tab</a></div> <figcaption><p>Haplotype networks for partial gene sequences of <em>Rickettsia bellii</em> and concatenated sequences. (A) atpA, (B) coxA, (C) gltA, (D) All genes concatenated. Red = North American R. <em>bellii</em>. Blue = South American <em>R. bellii</em>. Purple = Florida <em>R. bellii</em></p></figcaption></figure></section></section><section id="Sec11"><h2 class="pmc_sec_title">Discussion</h2> <p id="Par22">Our investigation aimed to determine whether tick infestation in cane toads and bacterial infection in ticks decreased from the core to the periphery of the cane toad’s distribution in Florida. We found evidence that cane toads at sites near the initial establishment were infested with <em>A. rotundatum</em>. Cane toads collected from Homestead, Miami, and West Palm Beach were infested with <em>A. rotundatum (</em>3/10 locations surveyed). These locations were along the south Atlantic coast of Florida where toad populations were introduced in the 1950’s and 1960’s (King and Krakauer <a href="#CR22" class="usa-link" aria-describedby="CR22">1966</a>; Krakauer <a href="#CR23" class="usa-link" aria-describedby="CR23">1968</a>; Wilson <a href="#CR57" class="usa-link" aria-describedby="CR57">2016</a>). Interestingly, cane toads from Fort Lauderdale lay between two infested populations yet this population did not have ticks, which exemplifies the stochastic nature of ectoparasite infestation among host populations. None of the cane toad populations > 120 km from the core of the introduction contained toads that were infested with ticks. In relation to the core population, these uninfested, peripheral populations lay to the west on Gulf Coast side of Florida or to the north in West Palm Beach. Cane toad populations in these locations did not become established until the early 2000’s and were geographically distant from the founder populations (Wilson <a href="#CR57" class="usa-link" aria-describedby="CR57">2016</a>). The lack of tick infestation along the invasion front is consistent with founder effects and provides support for the Enemy Release Hypothesis (Torchin et al. <a href="#CR54" class="usa-link" aria-describedby="CR54">2003</a>). Invasive species can establish populations without parasites via stochastic processes or lose their parasites as they advance along the expansion front due to low host densities or unfavorable environmental conditions for the invasive tick species (Torchin et al. <a href="#CR54" class="usa-link" aria-describedby="CR54">2003</a>; Phillips et al. <a href="#CR45" class="usa-link" aria-describedby="CR45">2010</a>).</p> <p id="Par23">Unfavorable environmental conditions for parasites at the periphery of an expanding distribution can further facilitate founder effects, but we did not find evidence for this additive effect. We hypothesized that because cane toads in Florida have been shown to prefer disturbed and urbanized habitats that may hinder tick survival (Meshaka et al. <a href="#CR37" class="usa-link" aria-describedby="CR37">2006</a>; Wilson <a href="#CR57" class="usa-link" aria-describedby="CR57">2016</a>), it could limit the expansion of this tick species on cane toads. <em>Amblyomma rotundatum</em> is a 3-host tick, meaning it must drop from its host after feeding, then molt and wait for another bloodmeal. The environment into which ticks drop is crucial to their survival because ticks spend more than 99% of their lifetime off-host (Needham <a href="#CR41" class="usa-link" aria-describedby="CR41">1991</a>, Diuk-Wasser et al. <a href="#CR12" class="usa-link" aria-describedby="CR12">2021</a>). Cane toads are commonly found in manmade ponds, canals, and residential yards (Meshaka et al. <a href="#CR37" class="usa-link" aria-describedby="CR37">2006</a>; Wilson <a href="#CR57" class="usa-link" aria-describedby="CR57">2016</a>, pers. Obs.), and these urban habitats may not have the appropriate microclimate to prevent tick desiccation (Needham <a href="#CR41" class="usa-link" aria-describedby="CR41">1991</a>). Urbanized habitats often lack preferred vegetation cover, moist soil, or leaf litter where ticks can take cover (Burtis et al. <a href="#CR6" class="usa-link" aria-describedby="CR6">2019</a>; Diuk-Wasser et al. <a href="#CR12" class="usa-link" aria-describedby="CR12">2021</a>). Studies in French Guiana (part of the native range of both cane toads and their ticks) have documented that urbanized habitat led to a loss of ticks on cane toads despite having sufficient toad populations to support tick transmission. In these cases, abiotic constraints in the urban environment prevented ticks from establishing (Devore et al. <a href="#CR11" class="usa-link" aria-describedby="CR11">2020</a>). Nonetheless, we found ticks in the heavily urbanized core population of cane toads that have been established for more than 60 years, suggesting that local environmental conditions were not the driving factor limiting expansion on cane toad populations.</p> <p id="Par24">The core to periphery gradient in tick infestation of cane toads did not support the hypothesis that this distribution was driven by spillback of <em>A. rotundatum</em> to toads. The widespread distribution of this tick on multiple reptile hosts sympatric to peripheral cane toad populations provided opportunity for spillback in the northern and western populations of cane toads, yet these populations were uninfested. Because the pattern of infestation was structured along a core to periphery gradient, it was unlikely a result of spillback of ticks from other established or native species. Indeed, cane toads were likely the initial hosts and one of the dispersal vectors of <em>A. rotundatum</em> that provided the mechanism for spillover of this tick to multiple native and non-native reptile species throughout Florida (Oliver et al. <a href="#CR42" class="usa-link" aria-describedby="CR42">1993</a>).</p> <p id="Par25">We found evidence of a likely exotic bacterial microbe throughout the tick populations that we sampled. We detected <em>R. bellii</em> in <em>A. rotundatum</em> at all three field sites where ticks were found. Thus, while tick dispersal among host toad populations may have been hindered, it appears that <em>Rickettsia</em> have successfully dispersed with their tick vector as predicted by their endosymbiont status and transovarial and transstadial transmission routes.</p> <p id="Par26">Previous investigations into the phylogenetics of <em>R. bellii</em> found that isolates collected from North America and South America formed separate clades and grouped together by tick genus and host preference (Krawczak et al. <a href="#CR24" class="usa-link" aria-describedby="CR24">2018</a>). In North America, <em>R. bellii</em> has primarily been reported in <em>Dermacentor</em> species (Krawczak et al. <a href="#CR24" class="usa-link" aria-describedby="CR24">2018</a>), while in South America it has been shown to infect ticks in the genera <em>Ixodes</em>, <em>Haemaphysalis</em> and <em>Amblyomma</em> (Krawczak et al. <a href="#CR24" class="usa-link" aria-describedby="CR24">2018</a>). Even though North American and South America <em>Rickettsia bellii</em> form distinct clades, there are few polymorphisms that differentiate isolates (< 0.5%) (Krawczak et al <a href="#CR24" class="usa-link" aria-describedby="CR24">2018</a>). We sequenced three rickettsial genes that each contained polymorphisms found in <em>R. bellii</em>. The atpA gene matched closely to <em>Rickettsia</em> sequences from <em>Amblyomma dissimile</em> in French Guiana<em>,</em> another avid reptile feeder in South America (Table <a href="#Tab3" class="usa-link">3</a>). Vector-pathogen affiliation and haplotype data from this atpA gene suggest <em>R. bellii</em> originated from South America. However, the coxA and gltA genes resembled strains recovered from different tick species and locations (Table <a href="#Tab3" class="usa-link">3</a>). This combination of gene sequence data could not clarify whether or not our <em>R. bellii</em> originated from South America. Thus, while the atpA gene is suggestive of a South American origin, further genomic analysis is needed to definitively determine the origin of <em>R. bellii</em> in Florida.</p> <p id="Par27">Studies of <em>Rickettsia</em> and toads thus far have only identified pathogens in ticks but not toad tissues (Luz et al. <a href="#CR35" class="usa-link" aria-describedby="CR35">2013</a>; Horta et al. <a href="#CR19" class="usa-link" aria-describedby="CR19">2006</a>). We did not detect <em>Rickettsia</em> species in attachment site or vent tissue collected from cane toads in our study. Tick attachment tissue examined in previous studies on cane toads in South America found that <em>A. rotundatum</em> bites can result in skin lesions (Luz et al. <a href="#CR35" class="usa-link" aria-describedby="CR35">2013</a>). Whether or not these lesions can harbor tick-borne pathogens is still unknown. Future studies should collect tissue from lesions and attachment sites and test them for tick-borne pathogens.</p></section><section id="Sec12"><h2 class="pmc_sec_title">Conclusions</h2> <p id="Par28">Overall, we found that <em>A. rotundatum</em> infests cane toads in areas < 120 km from the initial introduction sites in southern Florida, but not in populations further north or west. Cane toad populations in Homestead and Miami, sites at or near the initial introductions, continue to have tick infestations with <em>A. rotundatum.</em> The expansion of this tick species on cane toads into peripheral populations may have been restricted by demographic stochasticity associated with founder events rather than changes in habitat conditions from core to periphery. The core to periphery decline in infestation suggests that these processes are a more likely explanation than spill-back from established host populations because peripheral populations were sympatric with other reptiles hosting <em>A. rotundatum</em>. Our surveys found that <em>A. rotundatum</em> from cane toads were frequently infected with <em>Rickettsia bellii</em>, which was likely a South American strain that arrived in Florida with cane toads. This invasion system provides insight into how <em>Rickettsia</em> bacteria can be successfully transported during a host invasion and maintained during the expansion phase. Because many <em>Rickettsia</em> species are pathogenic, these findings suggest that pathogen pollution with exotic <em>Rickettsia</em> can impact human or animal welfare globally.</p></section><section id="Sec13"><h2 class="pmc_sec_title">Supplementary Information</h2> <p>Below is the link to the electronic supplementary material.</p> <section class="sm xbox font-sm" id="MOESM1"><div class="media p"><div class="caption"> <a href="/articles/instance/11213802/bin/10530_2024_3291_MOESM1_ESM.docx" data-ga-action="click_feat_suppl" class="usa-link">Supplementary file 1 (DOCX 16 kb)</a><sup> (16.2KB, docx) </sup> </div></div></section></section><section id="ack1" class="ack"><h2 class="pmc_sec_title">Acknowledgements</h2> <p>Special thanks to Yasmin Tavares, Chanakya Bhosale, Nathan Fox, and Natalie Pegg for assisting with cane toad collections. Thanks to Kylie Mendoza, Amber Hanna, Dylan Withee, Nathan Fox, and Samuel Scherneck for assisting with toad biopsies and data collection. Thanks for Savannah Cantrell for assisting with data entry.</p></section><section id="notes1"><h2 class="pmc_sec_title">Author contributions</h2> <p>CED coordinated the research project, collected toads, conducted pathogen testing, conducted phylogenetic analyses, wrote, and edited manuscript. MH coordinated the research project, collected toads, collected ticks, conducted toad biopsies, wrote, and edited the manuscript. AS assisted with toad biopsies and edited the manuscript. SB assisted with collection of cane toads and edited the manuscript. CMR supervised the research project, acquired funding, and edited the manuscript. SMW supervised the research project, acquired funding, and edited the manuscript. All authors have read and agreed to the published version of the manuscript.</p></section><section id="notes2"><h2 class="pmc_sec_title">Data availability</h2> <p>Sequence data with metadata have been deposited in NCBI Genbank.</p></section><section id="notes3"><h2 class="pmc_sec_title">Declarations</h2> <section id="FPar1"><h3 class="pmc_sec_title">Conflict of interest</h3> <p id="Par29">The authors declare no financial or non-financial conflicts of interest.</p></section><section id="FPar2"><h3 class="pmc_sec_title">Human and animal rights</h3> <p id="Par30">Collection methods for cane toads and ticks was approved by University of Florida Institutional Animal Care and Use Committee (#202111387).</p></section></section><section id="fn-group1" class="fn-group"><h2 class="pmc_sec_title">Footnotes</h2> <div class="fn-group p font-secondary-light font-sm"><div class="fn p" id="fn1"> <p><strong>Publisher's Note</strong></p> <p>Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.</p> </div></div></section><section id="Bib1" class="ref-list"><h2 class="pmc_sec_title">References</h2> <section id="Bib1_sec2"><ol class="ref-list font-sm"> <li id="CR1"> <cite>Acevedo AA, Lampo M, Cipriani R. The cane or marine toad, Rhinella marina (Anura, Bufonidae): two genetically and morphologically distinct species. Zootaxa. 2016;4103(6):574–586. doi: 10.11646/zootaxa.4103.6.7.</cite> [<a href="https://doi.org/10.11646/zootaxa.4103.6.7" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/27394759/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Zootaxa&title=The%20cane%20or%20marine%20toad,%20Rhinella%20marina%20(Anura,%20Bufonidae):%20two%20genetically%20and%20morphologically%20distinct%20species&author=AA%20Acevedo&author=M%20Lampo&author=R%20Cipriani&volume=4103&issue=6&publication_year=2016&pages=574-586&pmid=27394759&doi=10.11646/zootaxa.4103.6.7&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR2"><cite>American veterinary medical association. (2020). AVMA guidelines for the euthanasia of animals: 2020.0.1 edition. 121 pp</cite></li> <li id="CR3"> <cite>Barnett LK, Phillips BL, Heath ACG, Coates A, Hoskin CJ. The impact of parasites during range expansion of an invasive gecko. Parasitology. 2018;145(11):1400–1409. doi: 10.1017/S003118201800015X.</cite> [<a href="https://doi.org/10.1017/S003118201800015X" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/29441839/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Parasitology&title=The%20impact%20of%20parasites%20during%20range%20expansion%20of%20an%20invasive%20gecko&author=LK%20Barnett&author=BL%20Phillips&author=ACG%20Heath&author=A%20Coates&author=CJ%20Hoskin&volume=145&issue=11&publication_year=2018&pages=1400-1409&pmid=29441839&doi=10.1017/S003118201800015X&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR4"> <cite>Barré N, Uilenberg G. Spread of parasites transported with their hosts: case study of two species of cattle tick. Rev Sci Tech. 2010;29(1):149.</cite> [<a href="https://pubmed.ncbi.nlm.nih.gov/20617654/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Rev%20Sci%20Tech&title=Spread%20of%20parasites%20transported%20with%20their%20hosts:%20case%20study%20of%20two%20species%20of%20cattle%20tick&author=N%20Barr%C3%A9&author=G%20Uilenberg&volume=29&issue=1&publication_year=2010&pages=149&pmid=20617654&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR5"><cite>Burridge, MJ (2011) Non-native and invasive ticks. University Press of Florida</cite></li> <li id="CR6"> <cite>Burtis JC, Yavitt JB, Fahey TJ, Ostfeld RS. Ticks as soil-dwelling arthropods: an intersection between disease and soil ecology. J Med Entomol. 2019;56(6):1555–1564. doi: 10.1093/jme/tjz11.</cite> [<a href="https://doi.org/10.1093/jme/tjz11" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/31318035/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=J%20Med%20Entomol&title=Ticks%20as%20soil-dwelling%20arthropods:%20an%20intersection%20between%20disease%20and%20soil%20ecology&author=JC%20Burtis&author=JB%20Yavitt&author=TJ%20Fahey&author=RS%20Ostfeld&volume=56&issue=6&publication_year=2019&pages=1555-1564&pmid=31318035&doi=10.1093/jme/tjz11&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR7"> <cite>Capinha C, Seebens H, Cassey P, García-Díaz P, Lenzner B, Mang T, Moser D, Pyšek P, Rödder D, Scalera R, Winter M, Dullinger S, Essl F. Diversity, biogeography and the global flows of alien amphibians and reptiles. Divers Distrib. 2017;23(11):1313–1322. doi: 10.1111/ddi.12617.</cite> [<a href="https://doi.org/10.1111/ddi.12617" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Divers%20Distrib&title=Diversity,%20biogeography%20and%20the%20global%20flows%20of%20alien%20amphibians%20and%20reptiles&author=C%20Capinha&author=H%20Seebens&author=P%20Cassey&author=P%20Garc%C3%ADa-D%C3%ADaz&author=B%20Lenzner&volume=23&issue=11&publication_year=2017&pages=1313-1322&doi=10.1111/ddi.12617&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR8"> <cite>Chalkowski K, Lepczyk CA, Zohdy S. Parasite ecology of invasive species: conceptual framework and new hypotheses. Trends Parasitol. 2018;34(8):655–663. doi: 10.1016/j.pt.2018.05.008.</cite> [<a href="https://doi.org/10.1016/j.pt.2018.05.008" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/29935995/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Trends%20Parasitol&title=Parasite%20ecology%20of%20invasive%20species:%20conceptual%20framework%20and%20new%20hypotheses&author=K%20Chalkowski&author=CA%20Lepczyk&author=S%20Zohdy&volume=34&issue=8&publication_year=2018&pages=655-663&pmid=29935995&doi=10.1016/j.pt.2018.05.008&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR9"> <cite>Corn JL, Mertins JW, Hanson B, Snow S. First reports of ectoparasites collected from wild-caught exotic reptiles in Florida. J Med Entomol. 2014;48(1):94–100. doi: 10.1603/ME10065.</cite> [<a href="https://doi.org/10.1603/ME10065" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/21337954/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=J%20Med%20Entomol&title=First%20reports%20of%20ectoparasites%20collected%20from%20wild-caught%20exotic%20reptiles%20in%20Florida&author=JL%20Corn&author=JW%20Mertins&author=B%20Hanson&author=S%20Snow&volume=48&issue=1&publication_year=2014&pages=94-100&pmid=21337954&doi=10.1603/ME10065&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR10"> <cite>Cunningham AA, Daszak P, Rodriguez JP. Pathogen pollution: defining a parasitological threat to biodiversity conservation. J Parasitol. 2003;89(Suppl):S78–S83.</cite> [<a href="https://scholar.google.com/scholar_lookup?journal=J%20Parasitol&title=Pathogen%20pollution:%20defining%20a%20parasitological%20threat%20to%20biodiversity%20conservation&author=AA%20Cunningham&author=P%20Daszak&author=JP%20Rodriguez&volume=89&issue=Suppl&publication_year=2003&pages=S78-S83&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR11"> <cite>DeVore JL, Shine R, Ducatez S. Urbanization and translocation disrupt the relationship between host density and parasite abundance. J Anim Ecol. 2020;89(4):1122–1133. doi: 10.1111/1365-2656.13175.</cite> [<a href="https://doi.org/10.1111/1365-2656.13175" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/31945186/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=J%20Anim%20Ecol&title=Urbanization%20and%20translocation%20disrupt%20the%20relationship%20between%20host%20density%20and%20parasite%20abundance&author=JL%20DeVore&author=R%20Shine&author=S%20Ducatez&volume=89&issue=4&publication_year=2020&pages=1122-1133&pmid=31945186&doi=10.1111/1365-2656.13175&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR12"> <cite>Diuk-Wasser MA, VanAcker MC, Fernandez MP. Impact of land use changes and habitat fragmentation on the eco-epidemiology of tick-borne diseases. J Med Entomol. 2021;58(4):1546–1564. doi: 10.1093/jme/tjaa209.</cite> [<a href="https://doi.org/10.1093/jme/tjaa209" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/33095859/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=J%20Med%20Entomol&title=Impact%20of%20land%20use%20changes%20and%20habitat%20fragmentation%20on%20the%20eco-epidemiology%20of%20tick-borne%20diseases&author=MA%20Diuk-Wasser&author=MC%20VanAcker&author=MP%20Fernandez&volume=58&issue=4&publication_year=2021&pages=1546-1564&pmid=33095859&doi=10.1093/jme/tjaa209&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR201"><cite>EDDMapS. 2024. Early Detection & Distribution Mapping System. The University of Georgia - Center for Invasive Species and Ecosystem Health</cite></li> <li id="CR13"> <cite>El Karkouri K, Ghigo E, Raoult D, Fournier PE. Genomic evolution and adaptation of arthropod-associated Rickettsia. Sci Rep. 2022;12(1):1–15. doi: 10.1038/s41598-022-07725-z.</cite> [<a href="https://doi.org/10.1038/s41598-022-07725-z" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="/articles/PMC8907221/" class="usa-link">PMC free article</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/35264613/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Sci%20Rep&title=Genomic%20evolution%20and%20adaptation%20of%20arthropod-associated%20Rickettsia&author=K%20El%20Karkouri&author=E%20Ghigo&author=D%20Raoult&author=PE%20Fournier&volume=12&issue=1&publication_year=2022&pages=1-15&pmid=35264613&doi=10.1038/s41598-022-07725-z&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR14"> <cite>Ewen JG, Bensch S, Blackburn TM, Bonneaud C, Brown R, Cassey P, Clarke RH, Pérez-Tris J. Establishment of exotic parasites: the origins and characteristics of an avian malaria community in an isolated island avifauna. Ecol Lett. 2012;15(10):1112–1119. doi: 10.1111/j.1461-0248.2012.01833.x.</cite> [<a href="https://doi.org/10.1111/j.1461-0248.2012.01833.x" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/22788956/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Ecol%20Lett&title=Establishment%20of%20exotic%20parasites:%20the%20origins%20and%20characteristics%20of%20an%20avian%20malaria%20community%20in%20an%20isolated%20island%20avifauna&author=JG%20Ewen&author=S%20Bensch&author=TM%20Blackburn&author=C%20Bonneaud&author=R%20Brown&volume=15&issue=10&publication_year=2012&pages=1112-1119&pmid=22788956&doi=10.1111/j.1461-0248.2012.01833.x&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR15"> <cite>Floerl O, Inglis GJ, Dey K, Smith A. The importance of transport hubs in stepping-stone invasions. J Appl Ecol. 2009;46(1):37–45. doi: 10.1111/j.1365-2664.2008.01540.x.</cite> [<a href="https://doi.org/10.1111/j.1365-2664.2008.01540.x" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=J%20Appl%20Ecol&title=The%20importance%20of%20transport%20hubs%20in%20stepping-stone%20invasions&author=O%20Floerl&author=GJ%20Inglis&author=K%20Dey&author=A%20Smith&volume=46&issue=1&publication_year=2009&pages=37-45&doi=10.1111/j.1365-2664.2008.01540.x&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR16"> <cite>Guglielmone AA, Nava S. Hosts of Amblyomma dissimile Koch, 1844 and Amblyomma rotundatum Koch, 1844 (Acari: Ixodidae) Zootaxa. 2010 doi: 10.11646/ZOOTAXA.2541.1.2.</cite> [<a href="https://doi.org/10.11646/ZOOTAXA.2541.1.2" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Zootaxa&title=Hosts%20of%20Amblyomma%20dissimile%20Koch,%201844%20and%20Amblyomma%20rotundatum%20Koch,%201844%20(Acari:%20Ixodidae)&author=AA%20Guglielmone&author=S%20Nava&publication_year=2010&doi=10.11646/ZOOTAXA.2541.1.2&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR17"> <cite>Guzman-Cornejo C, Robbins RG, Guglielmone AA, Montiel-Parra G, Pérez TM. The Amblyomma (Acari: Ixodida: Ixodidae) of Mexico Identification keys, distribution and hosts. Zootaxa. 2011;2998(1):16–38. doi: 10.11646/ZOOTAXA.2998.1.2.</cite> [<a href="https://doi.org/10.11646/ZOOTAXA.2998.1.2" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Zootaxa&title=The%20Amblyomma%20(Acari:%20Ixodida:%20Ixodidae)%20of%20Mexico%20Identification%20keys,%20distribution%20and%20hosts&author=C%20Guzman-Cornejo&author=RG%20Robbins&author=AA%20Guglielmone&author=G%20Montiel-Parra&author=TM%20P%C3%A9rez&volume=2998&issue=1&publication_year=2011&pages=16-38&doi=10.11646/ZOOTAXA.2998.1.2&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR18"> <cite>Hanson BA, Frank PA, Mertins JW, Corn JL. Tick paralysis of a snake caused by Amblyomma rotundatum (Acari: Ixodidae) J Med Entomol. 2007;44(1):155–157. doi: 10.1093/jmedent/41.5.155.</cite> [<a href="https://doi.org/10.1093/jmedent/41.5.155" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/17294934/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=J%20Med%20Entomol&title=Tick%20paralysis%20of%20a%20snake%20caused%20by%20Amblyomma%20rotundatum%20(Acari:%20Ixodidae)&author=BA%20Hanson&author=PA%20Frank&author=JW%20Mertins&author=JL%20Corn&volume=44&issue=1&publication_year=2007&pages=155-157&pmid=17294934&doi=10.1093/jmedent/41.5.155&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR19"> <cite>Horta MC, Pinter A, Schumaker TTS, Labruna MB. Natural infection, transovarial transmission, and transstadial survival of Rickettsia bellii in the tick Ixodes loricatus (Acari: Ixodidae) from Brazil. Ann N Y Acad Sci. 2006;1078:285–290. doi: 10.1196/annals.1374.053.</cite> [<a href="https://doi.org/10.1196/annals.1374.053" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/17114723/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Ann%20N%20Y%20Acad%20Sci&title=Natural%20infection,%20transovarial%20transmission,%20and%20transstadial%20survival%20of%20Rickettsia%20bellii%20in%20the%20tick%20Ixodes%20loricatus%20(Acari:%20Ixodidae)%20from%20Brazil&author=MC%20Horta&author=A%20Pinter&author=TTS%20Schumaker&author=MB%20Labruna&volume=1078&publication_year=2006&pages=285-290&pmid=17114723&doi=10.1196/annals.1374.053&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR20"><cite>De Jesus, C.E. 2021. Surveillance and ecology of tick-borne pathogens and tick-host associations of reptiles and amphibians in Florida. Dissertation Thesis. University of Florida</cite></li> <li id="CR21"> <cite>Keirans JE, Oliver JH., Jr First description of the male and redescription of the immature stages of Amblyomma rotundatum (Acari: Ixodidae), a recently discovered tick in the U.S.A. J Parasitol. 1993;79(6):860–865. doi: 10.2307/3283722.</cite> [<a href="https://doi.org/10.2307/3283722" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/8277377/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=J%20Parasitol&title=First%20description%20of%20the%20male%20and%20redescription%20of%20the%20immature%20stages%20of%20Amblyomma%20rotundatum%20(Acari:%20Ixodidae),%20a%20recently%20discovered%20tick%20in%20the%20U.S.A.&author=JE%20Keirans&author=JH%20Oliver&volume=79&issue=6&publication_year=1993&pages=860-865&pmid=8277377&doi=10.2307/3283722&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR22"> <cite>King W, Krakauer T. The exotic herpetofauna of Southeast Florida. Q J Fla Acad Sci. 1966;29(2):144–154.</cite> [<a href="https://scholar.google.com/scholar_lookup?journal=Q%20J%20Fla%20Acad%20Sci&title=The%20exotic%20herpetofauna%20of%20Southeast%20Florida&author=W%20King&author=T%20Krakauer&volume=29&issue=2&publication_year=1966&pages=144-154&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR23"> <cite>Krakauer T. The ecology of the neotropical toad, Bufo marinus. South Fla Herpetol. 1968;24(3):214–221.</cite> [<a href="https://scholar.google.com/scholar_lookup?journal=South%20Fla%20Herpetol&title=The%20ecology%20of%20the%20neotropical%20toad,%20Bufo%20marinus&author=T%20Krakauer&volume=24&issue=3&publication_year=1968&pages=214-221&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR24"> <cite>Krawczak FS, Labruna MB, Hecht JA, Paddock CD, Karpathy SE. Genotypic characterization of Rickettsia bellii reveals distinct lineages in the United States and South America. Biomed Res Int. 2018;2018:8505483. doi: 10.1155/2018/8505483.</cite> [<a href="https://doi.org/10.1155/2018/8505483" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="/articles/PMC5911327/" class="usa-link">PMC free article</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/29850579/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Biomed%20Res%20Int&title=Genotypic%20characterization%20of%20Rickettsia%20bellii%20reveals%20distinct%20lineages%20in%20the%20United%20States%20and%20South%20America&author=FS%20Krawczak&author=MB%20Labruna&author=JA%20Hecht&author=CD%20Paddock&author=SE%20Karpathy&volume=2018&publication_year=2018&pages=8505483&pmid=29850579&doi=10.1155/2018/8505483&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR25"> <cite>Krysko KL, Burgess JP, Rochford MR, Gillette CR, CuevaD Enge KM, Somma LA, Stabile JL, Smith DC, Wasilewski JA, Others, Verified non-indigenous amphibians and reptiles in Florida from 1863 through 2010: outlining the invasion process and identifying invasion pathways and stages. Zootaxa. 2011;3028(1):1–64. doi: 10.11646/zootaxa.3028.1.1.</cite> [<a href="https://doi.org/10.11646/zootaxa.3028.1.1" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Zootaxa&title=Verified%20non-indigenous%20amphibians%20and%20reptiles%20in%20Florida%20from%201863%20through%202010:%20outlining%20the%20invasion%20process%20and%20identifying%20invasion%20pathways%20and%20stages&author=KL%20Krysko&author=JP%20Burgess&author=MR%20Rochford&author=CR%20Gillette&author=Enge%20KM%20CuevaD&volume=3028&issue=1&publication_year=2011&pages=1-64&doi=10.11646/zootaxa.3028.1.1&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR26"> <cite>Kumar S, Stecher G, Li M, Knyaz C, Tamura K. MEGA X: molecular evolutionary genetics analysis across computing platforms. Mol Biol Evol. 2018;35(6):1547–1549. doi: 10.1093/molbev/msy096.</cite> [<a href="https://doi.org/10.1093/molbev/msy096" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="/articles/PMC5967553/" class="usa-link">PMC free article</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/29722887/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Mol%20Biol%20Evol&title=MEGA%20X:%20molecular%20evolutionary%20genetics%20analysis%20across%20computing%20platforms&author=S%20Kumar&author=G%20Stecher&author=M%20Li&author=C%20Knyaz&author=K%20Tamura&volume=35&issue=6&publication_year=2018&pages=1547-1549&pmid=29722887&doi=10.1093/molbev/msy096&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR27"> <cite>Labruna MB. Ecology of Rickettsia in South America. Ann N Y Acad Sci. 2009;1166(1):156–166. doi: 10.1111/j.1749-6632.2009.04516.x.</cite> [<a href="https://doi.org/10.1111/j.1749-6632.2009.04516.x" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/19538276/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Ann%20N%20Y%20Acad%20Sci&title=Ecology%20of%20Rickettsia%20in%20South%20America&author=MB%20Labruna&volume=1166&issue=1&publication_year=2009&pages=156-166&pmid=19538276&doi=10.1111/j.1749-6632.2009.04516.x&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR28"> <cite>Labruna MB, Whitworth T, Bouyer DH, McBride J, Camargo LMA, Camargo EP, Popov V, Walker DH. Rickettsia bellii and Rickettsia amblyommii in Amblyomma ticks from the state of Rondônia, Western Amazon. Brazil J Med Entomol. 2004;41(6):1073–1081. doi: 10.1603/0022-2585-41.6.1073.</cite> [<a href="https://doi.org/10.1603/0022-2585-41.6.1073" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/15605647/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Brazil%20J%20Med%20Entomol&title=Rickettsia%20bellii%20and%20Rickettsia%20amblyommii%20in%20Amblyomma%20ticks%20from%20the%20state%20of%20Rond%C3%B4nia,%20Western%20Amazon&author=MB%20Labruna&author=T%20Whitworth&author=DH%20Bouyer&author=J%20McBride&author=LMA%20Camargo&volume=41&issue=6&publication_year=2004&pages=1073-1081&pmid=15605647&doi=10.1603/0022-2585-41.6.1073&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR29"> <cite>Laukaitis HJ, Macaluso KR. Unpacking the intricacies of Rickettsia–vector interactions. Trends Parasitol. 2021;37(8):734–746. doi: 10.1016/j.pt.2021.05.008.</cite> [<a href="https://doi.org/10.1016/j.pt.2021.05.008" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="/articles/PMC8344978/" class="usa-link">PMC free article</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/34162522/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Trends%20Parasitol&title=Unpacking%20the%20intricacies%20of%20Rickettsia%E2%80%93vector%20interactions&author=HJ%20Laukaitis&author=KR%20Macaluso&volume=37&issue=8&publication_year=2021&pages=734-746&pmid=34162522&doi=10.1016/j.pt.2021.05.008&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR30"> <cite>Leigh JW, Bryant D. Popart: full-feature software for haplotype network construction. Methods Ecol Evol. 2015;6(9):1110–1116. doi: 10.1111/2041-210x.12410.</cite> [<a href="https://doi.org/10.1111/2041-210x.12410" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Methods%20Ecol%20Evol&title=Popart:%20full-feature%20software%20for%20haplotype%20network%20construction&author=JW%20Leigh&author=D%20Bryant&volume=6&issue=9&publication_year=2015&pages=1110-1116&doi=10.1111/2041-210x.12410&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR31"><cite>Lever C (2001). <em>The cane toad: The history and ecology of a successful colonist</em>. Westbury Academic & Scientific Pub</cite></li> <li id="CR32"> <cite>Levin ML, Snellgrove AN, Zemtsova GE. Comparative value of blood and skin samples for diagnosis of spotted fever group rickettsial infection in model animals. Ticks Tick Borne Dis. 2016;7(5):1029–1034. doi: 10.1016/j.ttbdis.2016.05.011.</cite> [<a href="https://doi.org/10.1016/j.ttbdis.2016.05.011" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="/articles/PMC5661872/" class="usa-link">PMC free article</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/27282078/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Ticks%20Tick%20Borne%20Dis&title=Comparative%20value%20of%20blood%20and%20skin%20samples%20for%20diagnosis%20of%20spotted%20fever%20group%20rickettsial%20infection%20in%20model%20animals&author=ML%20Levin&author=AN%20Snellgrove&author=GE%20Zemtsova&volume=7&issue=5&publication_year=2016&pages=1029-1034&pmid=27282078&doi=10.1016/j.ttbdis.2016.05.011&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR33"> <cite>Lillywhite HB, Sheehy CM., III . The unique insular population of cottonmouth snakes at seahorse key. In: Lillywhite H, Martins M, editors. Islands and snakes: isolation and adaptive evolution. In Oxford University Press, New York: Oxford University Press. Islands and Snakes; 2019. p. 400.</cite> [<a href="https://scholar.google.com/scholar_lookup?title=Islands%20and%20snakes:%20isolation%20and%20adaptive%20evolution&author=HB%20Lillywhite&author=CM%20Sheehy&publication_year=2019&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR34"> <cite>Luong LT, Horn CJ, Brophy T. Mitey costly: energetic costs of parasite avoidance and infection. Physiol Biochem Zool. 2017;90(4):471–477. doi: 10.1086/691704471.</cite> [<a href="https://doi.org/10.1086/691704471" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/28414262/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Physiol%20Biochem%20Zool&title=Mitey%20costly:%20energetic%20costs%20of%20parasite%20avoidance%20and%20infection&author=LT%20Luong&author=CJ%20Horn&author=T%20Brophy&volume=90&issue=4&publication_year=2017&pages=471-477&pmid=28414262&doi=10.1086/691704471&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR35"> <cite>Luz HR, Faccini JLH, Pires MS, da Silva HR, Barros-Battesti DM. Life cycle and behavior of Amblyomma rotundatum (Acari: Ixodidae) under laboratory conditions and remarks on parasitism of toads in Brazil. Exp Appl Acarol. 2013;60(1):55–62. doi: 10.1007/s10493-012-9628-8.</cite> [<a href="https://doi.org/10.1007/s10493-012-9628-8" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/23100108/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Exp%20Appl%20Acarol&title=Life%20cycle%20and%20behavior%20of%20Amblyomma%20rotundatum%20(Acari:%20Ixodidae)%20under%20laboratory%20conditions%20and%20remarks%20on%20parasitism%20of%20toads%20in%20Brazil&author=HR%20Luz&author=JLH%20Faccini&author=MS%20Pires&author=HR%20da%20Silva&author=DM%20Barros-Battesti&volume=60&issue=1&publication_year=2013&pages=55-62&pmid=23100108&doi=10.1007/s10493-012-9628-8&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR36"> <cite>Luz HR, Silva-Santos E, Costa-Campos CE, Acosta I, Martins TF, Muñoz-Leal S, McIntosh D, Faccini JLH, Labruna MB. Detection of Rickettsia spp. in ticks parasitizing toads (Rhinella horribilis) in the northern Brazilian Amazon. Experimental and Applied Acarology. 2018;75(3):309–318. doi: 10.1007/s10493-018-0270-y.</cite> [<a href="https://doi.org/10.1007/s10493-018-0270-y" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/29846852/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Experimental%20and%20Applied%20Acarology&title=Detection%20of%20Rickettsia%20spp.%20in%20ticks%20parasitizing%20toads%20(Rhinella%20horribilis)%20in%20the%20northern%20Brazilian%20Amazon&author=HR%20Luz&author=E%20Silva-Santos&author=CE%20Costa-Campos&author=I%20Acosta&author=TF%20Martins&volume=75&issue=3&publication_year=2018&pages=309-318&pmid=29846852&doi=10.1007/s10493-018-0270-y&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR37"> <cite>Meshaka WE, DeVane J, Marshall SD. An island of cane toads (Bufo marinus) in an ocean of xeric uplands in south-central Florida. Fla Sci. 2006;69(3):169–176.</cite> [<a href="https://scholar.google.com/scholar_lookup?journal=Fla%20Sci&title=An%20island%20of%20cane%20toads%20(Bufo%20marinus)%20in%20an%20ocean%20of%20xeric%20uplands%20in%20south-central%20Florida&author=WE%20Meshaka&author=J%20DeVane&author=SD%20Marshall&volume=69&issue=3&publication_year=2006&pages=169-176&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR38"> <cite>Mittan-Moreau CS, Kelehear C, Toledo LF, Bacon J, Guayasamin JM, Snyder A, Zamudio KR. Cryptic lineages and standing genetic variation across independent cane toad introductions. Mol Ecol. 2022;31(24):6440–6656. doi: 10.1111/mec.16713.</cite> [<a href="https://doi.org/10.1111/mec.16713" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="/articles/PMC10091960/" class="usa-link">PMC free article</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/36198047/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Mol%20Ecol&title=Cryptic%20lineages%20and%20standing%20genetic%20variation%20across%20independent%20cane%20toad%20introductions&author=CS%20Mittan-Moreau&author=C%20Kelehear&author=LF%20Toledo&author=J%20Bacon&author=JM%20Guayasamin&volume=31&issue=24&publication_year=2022&pages=6440-6656&pmid=36198047&doi=10.1111/mec.16713&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR100"> <cite>Mittan CS, Zamudio KR (2019) Rapid adaptation to cold in the invasive cane toad <em>Rhinella marina</em>. Conservation Physiol 7(1):coy075. 10.1093/conphys/coy075</cite> [<a href="https://doi.org/10.1093/conphys/coy075" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="/articles/PMC6379050/" class="usa-link">PMC free article</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/30800317/" class="usa-link">PubMed</a>]</li> <li id="CR39"> <cite>Mooney HA, Cleland EE. The evolutionary impact of invasive species. Proc Natl Acad Sci U S A. 2001;98(10):5446–5451. doi: 10.1073/pnas.091093398.</cite> [<a href="https://doi.org/10.1073/pnas.091093398" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="/articles/PMC33232/" class="usa-link">PMC free article</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/11344292/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Proc%20Natl%20Acad%20Sci%20U%20S%20A&title=The%20evolutionary%20impact%20of%20invasive%20species&author=HA%20Mooney&author=EE%20Cleland&volume=98&issue=10&publication_year=2001&pages=5446-5451&pmid=11344292&doi=10.1073/pnas.091093398&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR40"> <cite>Nava S, Venzal JM, Acuña DG, Martins TF, Guglielmone AA. Ticks of the Southern Cone of America: diagnosis, distribution, and hosts with taxonomy. Ecology and Sanitary Importance: Academic Press; 2017. </cite> [<a href="https://scholar.google.com/scholar_lookup?title=Ticks%20of%20the%20Southern%20Cone%20of%20America:%20diagnosis,%20distribution,%20and%20hosts%20with%20taxonomy&author=S%20Nava&author=JM%20Venzal&author=DG%20Acu%C3%B1a&author=TF%20Martins&author=AA%20Guglielmone&publication_year=2017&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR41"> <cite>Needham GR, Teel PD. Off-host physiological ecology of ixodid ticks. Annu Rev Entomol. 1991;36:659–681. doi: 10.1146/annurev.en.36.010191.003303.</cite> [<a href="https://doi.org/10.1146/annurev.en.36.010191.003303" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/2006871/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Annu%20Rev%20Entomol&title=Off-host%20physiological%20ecology%20of%20ixodid%20ticks&author=GR%20Needham&author=PD%20Teel&volume=36&publication_year=1991&pages=659-681&pmid=2006871&doi=10.1146/annurev.en.36.010191.003303&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR42"> <cite>Oliver JH, Hayes MP, Keirans JE, Lavender DR. Establishment of the foreign parthenogenetic tick Amblyomma rotundatum (Acari: Ixodidae) in Florida. J Parasitol. 1993;79:786–786. doi: 10.2307/3283624.</cite> [<a href="https://doi.org/10.2307/3283624" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/8410557/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=J%20Parasitol&title=Establishment%20of%20the%20foreign%20parthenogenetic%20tick%20Amblyomma%20rotundatum%20(Acari:%20Ixodidae)%20in%20Florida&author=JH%20Oliver&author=MP%20Hayes&author=JE%20Keirans&author=DR%20Lavender&volume=79&publication_year=1993&pages=786-786&pmid=8410557&doi=10.2307/3283624&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR43"> <cite>Parola P, Paddock CD, Socolovschi C, Labruna MB, Mediannikov O, Kernif T, Abdad MY, Stenos J, Bitam I, Fournier PE, Raoult D. Update on tick-borne rickettsioses around the world: a geographic approach. Clin Microbiol Rev. 2013;26(4):657–702. doi: 10.1128/CMR.00032-13.</cite> [<a href="https://doi.org/10.1128/CMR.00032-13" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="/articles/PMC3811236/" class="usa-link">PMC free article</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/24092850/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Clin%20Microbiol%20Rev&title=Update%20on%20tick-borne%20rickettsioses%20around%20the%20world:%20a%20geographic%20approach&author=P%20Parola&author=CD%20Paddock&author=C%20Socolovschi&author=MB%20Labruna&author=O%20Mediannikov&volume=26&issue=4&publication_year=2013&pages=657-702&pmid=24092850&doi=10.1128/CMR.00032-13&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR45"> <cite>Phillips BL, Brown GP, Shine R. Life-history evolution in range-shifting populations. Ecology. 2010;91(6):1617–1627. doi: 10.1890/09-0910.1.</cite> [<a href="https://doi.org/10.1890/09-0910.1" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/20583704/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Ecology&title=Life-history%20evolution%20in%20range-shifting%20populations&author=BL%20Phillips&author=GP%20Brown&author=R%20Shine&volume=91&issue=6&publication_year=2010&pages=1617-1627&pmid=20583704&doi=10.1890/09-0910.1&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR46"> <cite>Piotrowski M, Rymaszewska A. Expansion of tick-borne rickettsioses in the world. Microorganisms. 2020;8(12):1906. doi: 10.3390/microorganisms8121906.</cite> [<a href="https://doi.org/10.3390/microorganisms8121906" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="/articles/PMC7760173/" class="usa-link">PMC free article</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/33266186/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Microorganisms&title=Expansion%20of%20tick-borne%20rickettsioses%20in%20the%20world&author=M%20Piotrowski&author=A%20Rymaszewska&volume=8&issue=12&publication_year=2020&pages=1906&pmid=33266186&doi=10.3390/microorganisms8121906&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR47"> <cite>Polo G, Luz HR, Regolin AL, Martins TF, Winck GR, da Silva HR, Onofrio VC, Labruna MB, Faccini JLH. Distribution modeling of Amblyomma rotundatum and Amblyomma dissimile in Brazil: estimates of environmental suitability. Parasitol Res. 2021;120(3):797–806. doi: 10.1007/s00436-020-06924-9.</cite> [<a href="https://doi.org/10.1007/s00436-020-06924-9" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/33068151/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Parasitol%20Res&title=Distribution%20modeling%20of%20Amblyomma%20rotundatum%20and%20Amblyomma%20dissimile%20in%20Brazil:%20estimates%20of%20environmental%20suitability&author=G%20Polo&author=HR%20Luz&author=AL%20Regolin&author=TF%20Martins&author=GR%20Winck&volume=120&issue=3&publication_year=2021&pages=797-806&pmid=33068151&doi=10.1007/s00436-020-06924-9&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR48"> <cite>Prenter J, Macneil C, Dick JTA, Dunn AM. Roles of parasites in animal invasions. Trends Ecol Evol. 2004;19(7):385–390. doi: 10.1016/j.tree.2004.05.002.</cite> [<a href="https://doi.org/10.1016/j.tree.2004.05.002" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/16701290/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Trends%20Ecol%20Evol&title=Roles%20of%20parasites%20in%20animal%20invasions&author=J%20Prenter&author=C%20Macneil&author=JTA%20Dick&author=AM%20Dunn&volume=19&issue=7&publication_year=2004&pages=385-390&pmid=16701290&doi=10.1016/j.tree.2004.05.002&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR49"><cite>Qiagen. (2014). <em>Gentra® Puregene® Handbook</em>.</cite></li> <li id="CR50"> <cite>Roux V, Rydkina E, Eremeeva M, Raoult D. Citrate synthase gene comparison, a new tool for phylogenetic analysis, and its application for the rickettsiae. Int J Syst Bacteriol. 1997;47(2):252–261. doi: 10.1099/00207713-47-2-252.</cite> [<a href="https://doi.org/10.1099/00207713-47-2-252" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/9103608/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Int%20J%20Syst%20Bacteriol&title=Citrate%20synthase%20gene%20comparison,%20a%20new%20tool%20for%20phylogenetic%20analysis,%20and%20its%20application%20for%20the%20rickettsiae&author=V%20Roux&author=E%20Rydkina&author=M%20Eremeeva&author=D%20Raoult&volume=47&issue=2&publication_year=1997&pages=252-261&pmid=9103608&doi=10.1099/00207713-47-2-252&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR200"><cite>Rubio AO, French CM Catenazzi A (2020) Morphological correlates of invasion in Florida cane toad (<em>Rhinella marina</em>) populations: Shortening of legs and reduction in leg asymmetry as populations become established. Acta oecologica 109:103652. 10.1016/j.actao.2020.103652</cite></li> <li id="CR51"> <cite>Sakai AK, Allendorf FW, Holt JS, Lodge DM, Molofsky J, With KA, Baughman S, Cabin RJ, Cohen JE, Ellstrand NC, McCauley DE, O’Neil P, Parker IM, Thompson JN, Weller SG. The population biology of invasive species. Annu Rev Ecol Syst. 2001;32(1):305–332. doi: 10.1146/annurev.ecolsys.32.081501.114037.</cite> [<a href="https://doi.org/10.1146/annurev.ecolsys.32.081501.114037" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Annu%20Rev%20Ecol%20Syst&title=The%20population%20biology%20of%20invasive%20species&author=AK%20Sakai&author=FW%20Allendorf&author=JS%20Holt&author=DM%20Lodge&author=J%20Molofsky&volume=32&issue=1&publication_year=2001&pages=305-332&doi=10.1146/annurev.ecolsys.32.081501.114037&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR52"> <cite>Sánchez-Montes S, Isaak-Delgado AB, Guzmán-Cornejo C, Rendón-Franco E, Muñoz-García CI, Bermúdez S, Morales-Diaz J, Cruz-Romero A, Romero-Salas D, Dzul-Rosado K, Lugo-Caballero C, Colunga-Salas P, Becker I. Rickettsia species in ticks that parasitize amphibians and reptiles: novel report from Mexico and review of the worldwide record. Ticks Tick-Borne Dis. 2019;10(5):987–994. doi: 10.1016/j.ttbdis.2019.04.013.</cite> [<a href="https://doi.org/10.1016/j.ttbdis.2019.04.013" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/31126748/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Ticks%20Tick-Borne%20Dis&title=Rickettsia%20species%20in%20ticks%20that%20parasitize%20amphibians%20and%20reptiles:%20novel%20report%20from%20Mexico%20and%20review%20of%20the%20worldwide%20record&author=S%20S%C3%A1nchez-Montes&author=AB%20Isaak-Delgado&author=C%20Guzm%C3%A1n-Cornejo&author=E%20Rend%C3%B3n-Franco&author=CI%20Mu%C3%B1oz-Garc%C3%ADa&volume=10&issue=5&publication_year=2019&pages=987-994&pmid=31126748&doi=10.1016/j.ttbdis.2019.04.013&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR53"> <cite>Thompson JD, Higgins DG, Gibson TJ. CLUSTAL W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids Res. 1994;22(22):4673–4680. doi: 10.1093/nar/22.22.4673.</cite> [<a href="https://doi.org/10.1093/nar/22.22.4673" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="/articles/PMC308517/" class="usa-link">PMC free article</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/7984417/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Nucleic%20Acids%20Res&title=CLUSTAL%20W:%20improving%20the%20sensitivity%20of%20progressive%20multiple%20sequence%20alignment%20through%20sequence%20weighting,%20position-specific%20gap%20penalties%20and%20weight%20matrix%20choice&author=JD%20Thompson&author=DG%20Higgins&author=TJ%20Gibson&volume=22&issue=22&publication_year=1994&pages=4673-4680&pmid=7984417&doi=10.1093/nar/22.22.4673&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR54"> <cite>Torchin ME, Lafferty KD, Dobson AP, McKenzie VJ, Kuris AM. Introduced species and their missing parasites. Nature. 2003;421(6923):628–630. doi: 10.1038/nature01346.</cite> [<a href="https://doi.org/10.1038/nature01346" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/12571595/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Nature&title=Introduced%20species%20and%20their%20missing%20parasites&author=ME%20Torchin&author=KD%20Lafferty&author=AP%20Dobson&author=VJ%20McKenzie&author=AM%20Kuris&volume=421&issue=6923&publication_year=2003&pages=628-630&pmid=12571595&doi=10.1038/nature01346&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR55"> <cite>van Kleunen M, Dawson W, Maurel N. Characteristics of successful alien plants. Mol Ecol. 2015;24(9):1954–1968. doi: 10.1002/9781119072799.ch3.</cite> [<a href="https://doi.org/10.1002/9781119072799.ch3" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/25421056/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=Mol%20Ecol&title=Characteristics%20of%20successful%20alien%20plants&author=M%20van%20Kleunen&author=W%20Dawson&author=N%20Maurel&volume=24&issue=9&publication_year=2015&pages=1954-1968&pmid=25421056&doi=10.1002/9781119072799.ch3&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR56"> <cite>Weinert LA, Werren JH, Aebi A, Stone GN, Jiggins FM. Evolution and diversity of Rickettsia bacteria. BMC Biol. 2009;7(1):1–15. doi: 10.1186/1741-7007-7-6.</cite> [<a href="https://doi.org/10.1186/1741-7007-7-6" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">DOI</a>] [<a href="/articles/PMC2662801/" class="usa-link">PMC free article</a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/19187530/" class="usa-link">PubMed</a>] [<a href="https://scholar.google.com/scholar_lookup?journal=BMC%20Biol&title=Evolution%20and%20diversity%20of%20Rickettsia%20bacteria&author=LA%20Weinert&author=JH%20Werren&author=A%20Aebi&author=GN%20Stone&author=FM%20Jiggins&volume=7&issue=1&publication_year=2009&pages=1-15&pmid=19187530&doi=10.1186/1741-7007-7-6&" class="usa-link usa-link--external" data-ga-action="click_feat_suppl" target="_blank" rel="noopener noreferrer">Google Scholar</a>]</li> <li id="CR57"><cite>Wilson, A. C. (2016). Distribution of cane toads (<em>Rhinella horribilis</em>) in Florida and their status in natural areas. PhD Thesis. University of Florida</cite></li> </ol></section></section><section id="_ad93_" lang="en" class="associated-data"><h2 class="pmc_sec_title">Associated Data</h2> <p class="font-secondary"><em>This section collects any data citations, data availability statements, or supplementary materials included in this article.</em></p> <section id="_adsm93_" lang="en" class="supplementary-materials"><h3 class="pmc_sec_title">Supplementary Materials</h3> <section class="sm xbox font-sm" id="db_ds_supplementary-material1_reqid_"><div class="media p"><div class="caption"> <a href="/articles/instance/11213802/bin/10530_2024_3291_MOESM1_ESM.docx" data-ga-action="click_feat_suppl" class="usa-link">Supplementary file 1 (DOCX 16 kb)</a><sup> (16.2KB, docx) </sup> </div></div></section></section><section id="_adda93_" lang="en" class="data-availability-statement"><h3 class="pmc_sec_title">Data Availability Statement</h3> <p>Sequence data with metadata have been deposited in NCBI Genbank.</p></section></section></section><footer class="p courtesy-note font-secondary font-sm text-center"><hr class="headless"> <p>Articles from Biological Invasions are provided here courtesy of <strong>Springer</strong></p></footer></section></article> </main> </div> </div> </div>  <div class="pmc-sidenav desktop:grid-col-4 display-flex"> <section class="pmc-sidenav__container" aria-label="Article resources and navigation"> <button type="button" class="usa-button pmc-sidenav__container__close usa-button--unstyled"> <img src="/static/img/usa-icons/close.svg" role="img" alt="Close" /> </button> <div class="display-none desktop:display-block"> <section class="margin-top-4 desktop:margin-top-0"> <h2 class="margin-top-0">ACTIONS</h2> <ul class="usa-list usa-list--unstyled usa-list--actions"> <li> <a href="https://doi.org/10.1007/s10530-024-03291-9" class="usa-button usa-button--outline width-24 font-xs usa-link--external padding-left-0 padding-right-0" target="_blank" rel="noreferrer noopener" data-ga-category="actions" data-ga-action="click" data-ga-label="publisher_link_desktop" > <span class="height-3 display-inline-flex flex-align-center">View on publisher site</span> </a> </li> <li> <a href="pdf/10530_2024_Article_3291.pdf" class="usa-button usa-button--outline width-24 display-inline-flex flex-align-center flex-justify-start padding-left-1" data-ga-category="actions" data-ga-action="click" data-ga-label="pdf_download_desktop" > <svg class="usa-icon width-3 height-3" aria-hidden="true" focusable="false" role="img" hidden> <use xlink:href="/static/img/sprite.svg#file_download"></use> </svg> <span class="display-inline-flex flex-justify-center flex-1">PDF (974.8 KB)</span> </a> </li> <li> <button role="button" class="usa-button width-24 citation-dialog-trigger display-inline-flex flex-align-center flex-justify-start padding-left-1" aria-label="Open dialog with citation text in different styles" data-ga-category="actions" data-ga-action="open" data-ga-label="cite_desktop" data-all-citations-url="/resources/citations/11213802/" data-citation-style="nlm" data-download-format-link="/resources/citations/11213802/export/" > <svg class="usa-icon width-3 height-3" aria-hidden="true" focusable="false" role="img" hidden> <use xlink:href="/static/img/sprite.svg#format_quote"></use> </svg> <span class="display-inline-flex flex-justify-center flex-1 button-label">Cite</span> </button> </li> <li> <button class="usa-button width-24 collections-dialog-trigger collections-button display-inline-flex flex-align-center flex-justify-start padding-left-1 collections-button-empty" aria-label="Save article in MyNCBI collections." data-ga-category="actions" data-ga-action="click" data-ga-label="collections_button_desktop" data-collections-open-dialog-enabled="false" data-collections-open-dialog-url="https://account.ncbi.nlm.nih.gov/?back_url=https%3A%2F%2Fpmc.ncbi.nlm.nih.gov%2Farticles%2FPMC11213802%2F%23open-collections-dialog" data-in-collections="false"> <svg class="usa-icon width-3 height-3 usa-icon--bookmark-full" aria-hidden="true" focusable="false" role="img" hidden> <use xlink:href="/static/img/action-bookmark-full.svg#icon"></use> </svg> <svg class="usa-icon width-3 height-3 usa-icon--bookmark-empty" aria-hidden="true" focusable="false" role="img" hidden> <use xlink:href="/static/img/action-bookmark-empty.svg#icon"></use> </svg> <span class="display-inline-flex flex-justify-center flex-1">Collections</span> </button> </li> <li class="pmc-permalink"> <button type="button" class="usa-button usa-button--outline width-24 display-inline-flex flex-align-center flex-justify padding-left-1 shadow-none" aria-label="Show article permalink" aria-expanded="false" aria-haspopup="true" data-ga-category="actions" data-ga-action="open" data-ga-label="permalink_desktop" > <svg class="usa-icon width-3 height-3" aria-hidden="true" focusable="false" role="img" hidden> <use xlink:href="/static/img/sprite.svg#share"></use> </svg> <span class="display-inline-flex flex-justify-center flex-1 button-label">Permalink</span> </button> <div class="pmc-permalink__dropdown" hidden> <div class="pmc-permalink__dropdown__container"> <h2 class="usa-modal__heading margin-top-0 margin-bottom-2">PERMALINK</h2> <div class="pmc-permalink__dropdown__content"> <input type="text" class="usa-input" value="https://pmc.ncbi.nlm.nih.gov/articles/PMC11213802/" aria-label="Article permalink"> <button class="usa-button display-inline-flex pmc-permalink__dropdown__copy__btn margin-right-0" title="Copy article permalink" data-ga-category="save_share" data-ga-action="link" data-ga-label="copy_link"> <svg class="usa-icon" aria-hidden="true" focusable="false" role="img"> <use xlink:href="/static/img/sprite.svg#content_copy"></use> </svg> <span class="margin-left-1">Copy</span> </button> </div> </div> </div> </li> </ul> </section> </div> <section class="pmc-resources margin-top-6 desktop:margin-top-4" data-page-path="/articles/PMC11213802/"> <h2 class="margin-top-0">RESOURCES</h2> <div class="usa-accordion usa-accordion--multiselectable" data-allow-multiple> <h3 class="usa-accordion__heading"> <button type="button" class="usa-accordion__button" aria-expanded="false" aria-controls="resources-similar-articles" data-ga-category="resources_accordion" data-ga-action="open_similar_articles" data-ga-label="/articles/PMC11213802/" data-action-open="open_similar_articles" data-action-close="close_similar_articles" > Similar articles </button> </h3> <div id="resources-similar-articles" class="usa-accordion__content usa-prose" data-source-url="/resources/similar-article-links/38947421/" > </div> <h3 class="usa-accordion__heading"> <button type="button" class="usa-accordion__button" aria-expanded="false" aria-controls="resources-cited-by-other-articles" data-ga-category="resources_accordion" data-ga-action="open_cited_by" data-ga-label="/articles/PMC11213802/" data-action-open="open_cited_by" data-action-close="close_cited_by" > Cited by other articles </button> </h3> <div id="resources-cited-by-other-articles" class="usa-accordion__content usa-prose" data-source-url="/resources/cited-by-links/38947421/" > </div> <h3 class="usa-accordion__heading"> <button type="button" class="usa-accordion__button" aria-expanded="false" aria-controls="resources-links-to-ncbi-databases" data-ga-category="resources_accordion" data-ga-action="open_NCBI_links" data-ga-label="/articles/PMC11213802/" data-action-open="open_NCBI_links" data-action-close="close_NCBI_link" > Links to NCBI Databases </button> </h3> <div id="resources-links-to-ncbi-databases" class="usa-accordion__content usa-prose" data-source-url="/resources/db-links/11213802/" > </div> </div> </section> <section class="usa-in-page-nav usa-in-page-nav--wide margin-top-6 desktop:margin-top-4" data-title-text="On this page" data-title-heading-level="h2" data-scroll-offset="0" data-root-margin="-10% 0px -80% 0px" data-main-content-selector="main" data-threshold="1" hidden ></section> </section> </div> <div class="overlay" role="dialog" aria-label="Citation Dialog" hidden> <div class="dialog citation-dialog" aria-hidden="true"> <div class="display-inline-flex flex-align-center flex-justify width-full margin-bottom-2"> <h2 class="usa-modal__heading margin-0">Cite</h2> <button type="button" class="usa-button usa-button--unstyled close-overlay text-black width-auto" tabindex="1"> <svg class="usa-icon width-3 height-3" aria-hidden="true" focusable="false" role="img"> <use xlink:href="/static/img/sprite.svg#close"></use> </svg> </button> </div> <div class="citation-text-block"> <div class="citation-text margin-bottom-2"></div> <ul class="usa-list usa-list--unstyled display-inline-flex flex-justify width-full flex-align-center"> <li> <button class="usa-button usa-button--unstyled text-no-underline display-flex flex-align-center copy-button dialog-focus" data-ga-category="save_share" data-ga-action="cite" data-ga-label="copy" tabindex="2"> <svg class="usa-icon width-3 height-3" aria-hidden="true" focusable="false" role="img"> <use xlink:href="/static/img/sprite.svg#content_copy"></use> </svg> <span>Copy</span> </button> </li> <li> <a href="#" role="button" class="usa-button usa-button--unstyled text-no-underline display-flex flex-align-center export-button" data-ga-category="save_share" data-ga-action="cite" data-ga-label="download" title="Download a file for external citation management software" tabindex="3"> <svg class="usa-icon width-3 height-3" aria-hidden="true" focusable="false" role="img"> <use xlink:href="/static/img/sprite.svg#file_download"></use> </svg> <span class="display-none mobile-lg:display-inline">Download .nbib</span> <span class="display-inline mobile-lg:display-none">.nbib</span> </a> </li> <li> <div class="display-inline-flex flex-align-center"> <label class="usa-label margin-top-0">Format:</label> <select aria-label="Format" class="usa-select citation-style-selector padding-1 margin-top-0 border-0 padding-right-4" tabindex="4" > <option data-style-url-name="ama" value="AMA" > AMA </option> <option data-style-url-name="apa" value="APA" > APA </option> <option data-style-url-name="mla" value="MLA" > MLA </option> <option data-style-url-name="nlm" value="NLM" selected="selected"> NLM </option> </select> </div> </li> </ul> </div> </div> </div> <div class="overlay" role="dialog" hidden> <div id="collections-action-dialog" class="dialog collections-dialog" aria-hidden="true"> <div class="display-inline-flex flex-align-center flex-justify width-full margin-bottom-2"> <h2 class="usa-modal__heading margin-0">Add to Collections</h2> </div> <div class="collections-action-panel action-panel"> <form id="collections-action-dialog-form" class="usa-form maxw-full collections-action-panel-form action-panel-content action-form action-panel-smaller-selectors" data-existing-collections-url="/list-existing-collections/" data-add-to-existing-collection-url="/add-to-existing-collection/" data-create-and-add-to-new-collection-url="/create-and-add-to-new-collection/" data-myncbi-max-collection-name-length="100" data-collections-root-url="https://www.ncbi.nlm.nih.gov/myncbi/collections/"> <input type="hidden" name="csrfmiddlewaretoken" value="urWB1H943iCd3hcU4tjaWUa5PLGHfo1lWVZEMM81neVOBEhTZJwTyvcsMh54oal3"> <fieldset class="usa-fieldset margin-bottom-2"> <div class="usa-radio"> <input type="radio" id="collections-action-dialog-new" class="usa-radio__input usa-radio__input--tile collections-new margin-top-0" name="collections" value="new" data-ga-category="collections_button" data-ga-action="click" data-ga-label="collections_radio_new" /> <label class="usa-radio__label" for="collections-action-dialog-new">Create a new collection</label> </div> <div class="usa-radio"> <input type="radio" id="collections-action-dialog-existing" class="usa-radio__input usa-radio__input--tile collections-existing" name="collections" value="existing" checked="true" data-ga-category="collections_button" data-ga-action="click" data-ga-label="collections_radio_existing" /> <label class="usa-radio__label" for="collections-action-dialog-existing">Add to an existing collection</label> </div> </fieldset> <fieldset class="usa-fieldset margin-bottom-2"> <div class="action-panel-control-wrap new-collections-controls"> <label for="collections-action-dialog-add-to-new" class="usa-label margin-top-0"> Name your collection <abbr title="required" class="usa-hint usa-hint--required text-no-underline">*</abbr> </label> <input type="text" name="add-to-new-collection" id="collections-action-dialog-add-to-new" class="usa-input collections-action-add-to-new" pattern="[^"&=<>/]*" title="The following characters are not allowed in the Name field: "&=<>/" maxlength="" data-ga-category="collections_button" data-ga-action="create_collection" data-ga-label="non_favorties_collection" required /> </div> <div class="action-panel-control-wrap existing-collections-controls"> <label for="collections-action-dialog-add-to-existing" class="usa-label margin-top-0"> Choose a collection </label> <select id="collections-action-dialog-add-to-existing" class="usa-select collections-action-add-to-existing" data-ga-category="collections_button" data-ga-action="select_collection" data-ga-label="($('.collections-action-add-to-existing').val() === 'Favorites') ? 'Favorites' : 'non_favorites_collection'"> </select> <div class="collections-retry-load-on-error usa-input-error-message selection-validation-message"> Unable to load your collection due to an error<br> <a href="#">Please try again</a> </div> </div> </fieldset> <div class="display-inline-flex"> <button class="usa-button margin-top-0 action-panel-submit" type="submit" data-loading-label="Adding..." data-pinger-ignore data-ga-category="collections_button" data-ga-action="click" data-ga-label="add"> Add </button> <button class="usa-button usa-button--outline margin-top-0 action-panel-cancel" aria-label="Close 'Add to Collections' panel" ref="linksrc=close_collections_panel" data-ga-category="collections_button" data-ga-action="click" data-ga-label="cancel"> Cancel </button> </div> </form> </div> </div> </div> </div> </div> </div> <footer class="ncbi-footer ncbi-dark-background " > <div class="ncbi-footer__icon-section"> <div class="ncbi-footer__social-header"> Follow NCBI </div> <div class="grid-container ncbi-footer__ncbi-social-icons-container"> <a href="https://twitter.com/ncbi" class="ncbi-footer__social-icon ncbi-footer__social-icon--gray" target="_blank" rel="noreferrer noopener"> <svg width="40" height="40" viewBox="0 0 40 40" fill="none" xmlns="http://www.w3.org/2000/svg" focusable="false" aria-hidden="true"> <path d="m6.067 8 10.81 13.9L6 33.2h4.2l8.4-9.1 7.068 9.1H34L22.8 18.5 31.9 8h-3.5l-7.7 8.4L14.401 8H6.067Zm3.6 1.734h3.266l16.8 21.732H26.57L9.668 9.734Z"> </path> </svg> <span class="usa-sr-only">NCBI on X (formerly known as Twitter)</span> </a> <a href="https://www.facebook.com/ncbi.nlm" class="ncbi-footer__social-icon ncbi-footer__social-icon--gray" target="_blank" rel="noreferrer noopener"> <svg width="16" height="29" focusable="false" aria-hidden="true" viewBox="0 0 16 29" fill="none" xmlns="http://www.w3.org/2000/svg"> <path d="M3.8809 21.4002C3.8809 19.0932 3.8809 16.7876 3.8809 14.478C3.8809 14.2117 3.80103 14.1452 3.54278 14.1492C2.53372 14.1638 1.52334 14.1492 0.514288 14.1598C0.302626 14.1598 0.248047 14.0972 0.248047 13.8936C0.256034 12.4585 0.256034 11.0239 0.248047 9.58978C0.248047 9.37013 0.302626 9.30224 0.528931 9.3049C1.53798 9.31688 2.54837 9.3049 3.55742 9.31555C3.80103 9.31555 3.8809 9.26097 3.87957 9.00272C3.87158 8.00565 3.85428 7.00592 3.90753 6.00884C3.97142 4.83339 4.31487 3.73115 5.04437 2.78467C5.93095 1.63318 7.15699 1.09005 8.56141 0.967577C10.5582 0.79319 12.555 0.982221 14.5518 0.927641C14.7102 0.927641 14.7462 0.99287 14.7449 1.13664C14.7449 2.581 14.7449 4.02668 14.7449 5.47104C14.7449 5.67604 14.6517 5.68669 14.4946 5.68669C13.4523 5.68669 12.4113 5.68669 11.3703 5.68669C10.3506 5.68669 9.92057 6.10868 9.90593 7.13904C9.89661 7.7647 9.91525 8.39303 9.89794 9.01869C9.88995 9.26364 9.96583 9.31822 10.2015 9.31688C11.7204 9.30623 13.2393 9.31688 14.7595 9.3049C15.0257 9.3049 15.0723 9.3728 15.0444 9.62439C14.89 10.9849 14.7515 12.3467 14.6144 13.7085C14.5691 14.1571 14.5785 14.1585 14.1458 14.1585C12.8386 14.1585 11.5313 14.1665 10.2254 14.1518C9.95119 14.1518 9.89794 14.2317 9.89794 14.4899C9.90593 19.0799 9.89794 23.6752 9.91125 28.2612C9.91125 28.5674 9.8407 28.646 9.53186 28.6433C7.77866 28.6273 6.02414 28.6366 4.27094 28.634C3.82499 28.634 3.87158 28.6992 3.87158 28.22C3.87602 25.9472 3.87913 23.6739 3.8809 21.4002Z"> </path> </svg> <span class="usa-sr-only">NCBI on Facebook</span> </a> <a href="https://www.linkedin.com/company/ncbinlm" class="ncbi-footer__social-icon ncbi-footer__social-icon--gray" target="_blank" rel="noreferrer noopener"> <svg width="25" height="23" viewBox="0 0 26 24" fill="none" xmlns="http://www.w3.org/2000/svg" focusable="false" aria-hidden="true"> <path d="M14.6983 9.98423C15.6302 9.24808 16.5926 8.74754 17.6762 8.51991C19.673 8.09126 21.554 8.30824 23.1262 9.7526C24.2351 10.7723 24.7529 12.1115 25.0165 13.5612C25.1486 14.3363 25.2105 15.1218 25.2015 15.9081C25.2015 18.3043 25.2015 20.6898 25.2082 23.0806C25.2082 23.3468 25.1549 23.444 24.8621 23.4414C23.1297 23.4272 21.3992 23.4272 19.6704 23.4414C19.4041 23.4414 19.3429 23.3588 19.3442 23.1019C19.3535 20.5194 19.3442 17.9368 19.3442 15.3543C19.3442 14.0005 18.3258 12.9448 17.0266 12.9488C15.7273 12.9528 14.6983 14.0071 14.6983 15.361C14.6983 17.9328 14.6917 20.5047 14.6983 23.0753C14.6983 23.3708 14.6198 23.444 14.3296 23.4427C12.6185 23.4294 10.9079 23.4294 9.19779 23.4427C8.93155 23.4427 8.86099 23.3735 8.86232 23.1086C8.8783 19.7619 8.88628 16.4144 8.88628 13.066C8.88628 11.5688 8.87874 10.0708 8.86365 8.57182C8.86365 8.3575 8.90758 8.27896 9.14054 8.28029C10.9048 8.29094 12.6687 8.29094 14.4321 8.28029C14.6464 8.28029 14.6983 8.34818 14.6983 8.54653C14.6903 9.00047 14.6983 9.45441 14.6983 9.98423Z"> </path> <path d="M6.55316 15.8443C6.55316 18.2564 6.55316 20.6699 6.55316 23.082C6.55316 23.3629 6.48127 23.4388 6.19906 23.4374C4.47737 23.4241 2.75568 23.4241 1.03399 23.4374C0.767751 23.4374 0.69986 23.3629 0.701191 23.1006C0.709178 18.2648 0.709178 13.4281 0.701191 8.59053C0.701191 8.34026 0.765089 8.27237 1.01669 8.2737C2.74991 8.28435 4.48048 8.28435 6.20838 8.2737C6.47462 8.2737 6.5465 8.33627 6.54517 8.6065C6.54783 11.0186 6.55316 13.4308 6.55316 15.8443Z"> </path> <path d="M3.65878 0.243898C5.36804 0.243898 6.58743 1.45529 6.58743 3.1406C6.58743 4.75801 5.32145 5.95742 3.60819 5.96807C3.22177 5.97614 2.83768 5.90639 2.47877 5.76299C2.11985 5.61959 1.79344 5.40546 1.51897 5.13334C1.24449 4.86123 1.02755 4.53668 0.881058 4.17902C0.734563 3.82136 0.661505 3.43788 0.666231 3.05141C0.67555 1.42601 1.9362 0.242566 3.65878 0.243898Z"> </path> </svg> <span class="usa-sr-only">NCBI on LinkedIn</span> </a> <a href="https://github.com/ncbi" class="ncbi-footer__social-icon ncbi-footer__social-icon--gray" target="_blank" rel="noreferrer noopener"> <svg width="28" height="27" viewBox="0 0 28 28" fill="none" xmlns="http://www.w3.org/2000/svg" focusable="false" aria-hidden="true"> <path d="M16.7228 20.6334C17.5057 20.5527 18.2786 20.3944 19.0301 20.1608C21.3108 19.4193 22.5822 17.8259 22.963 15.4909C23.1228 14.5112 23.1814 13.5287 22.9883 12.5437C22.8106 11.6423 22.4013 10.8028 21.8007 10.1076C21.7526 10.0605 21.7197 10 21.7064 9.934C21.6931 9.86799 21.7 9.79952 21.7262 9.73748C22.0856 8.6206 21.9711 7.51969 21.601 6.42677C21.582 6.3497 21.5345 6.2827 21.468 6.23923C21.4016 6.19577 21.3211 6.17906 21.2429 6.19248C20.7329 6.21649 20.2313 6.33051 19.7611 6.52928C19.1103 6.7908 18.4899 7.12198 17.9104 7.51703C17.84 7.56996 17.7581 7.60551 17.6713 7.62078C17.5846 7.63605 17.4954 7.6306 17.4112 7.60489C15.2596 7.05882 13.0054 7.06203 10.8554 7.61421C10.7806 7.63586 10.7018 7.63967 10.6253 7.62534C10.5487 7.611 10.4766 7.57892 10.4148 7.53167C9.64788 7.03247 8.85171 6.58918 7.96368 6.33359C7.65781 6.24338 7.34123 6.19458 7.02239 6.18849C6.94879 6.17986 6.87462 6.19893 6.81432 6.242C6.75402 6.28507 6.71191 6.34904 6.69621 6.42145C6.32342 7.51437 6.2209 8.61527 6.56307 9.73348C6.59635 9.84264 6.64694 9.93316 6.54177 10.0516C5.47666 11.2604 5.09988 12.6834 5.19574 14.2676C5.2663 15.4244 5.46201 16.5466 6.01454 17.5769C6.84399 19.1171 8.21664 19.9119 9.85158 20.3352C10.3938 20.4706 10.9444 20.5698 11.4998 20.632C11.5384 20.7492 11.4506 20.7798 11.408 20.8291C11.1734 21.1179 10.9894 21.4441 10.8634 21.7942C10.7622 22.0458 10.8315 22.4039 10.6065 22.5516C10.263 22.7766 9.83827 22.8485 9.42421 22.8871C8.17936 23.0056 7.26471 22.4877 6.6283 21.4348C6.25552 20.8184 5.76956 20.3325 5.08523 20.0663C4.76981 19.9325 4.42139 19.8967 4.08537 19.9638C3.7898 20.029 3.73788 20.1901 3.93891 20.4111C4.03639 20.5234 4.14989 20.6207 4.27575 20.6999C4.9796 21.1318 5.51717 21.7884 5.80152 22.5636C6.37002 23.9973 7.48039 24.5697 8.93825 24.6323C9.43741 24.6575 9.93768 24.615 10.4254 24.5058C10.5892 24.4672 10.6531 24.4872 10.6517 24.6762C10.6451 25.4936 10.6637 26.3123 10.6517 27.131C10.6517 27.6635 10.1684 27.9297 9.58663 27.7393C8.17396 27.2671 6.84977 26.5631 5.66838 25.656C2.59555 23.2891 0.720966 20.1861 0.217704 16.3376C-0.357453 11.9127 0.911353 8.00824 3.98551 4.73881C6.11909 2.42656 8.99932 0.939975 12.1203 0.540191C16.5351 -0.0601815 20.4347 1.14323 23.7232 4.16373C26.2449 6.47869 27.724 9.37672 28.1048 12.7726C28.5828 17.0325 27.3686 20.7945 24.4768 23.9827C22.9762 25.6323 21.0956 26.8908 18.9982 27.6488C18.8783 27.6927 18.7585 27.738 18.636 27.7726C18.0356 27.9404 17.6189 27.6395 17.6189 27.0098C17.6189 25.7452 17.6308 24.4806 17.6295 23.2159C17.6329 22.9506 17.6128 22.6856 17.5696 22.4238C17.4325 21.6664 17.3419 21.484 16.7228 20.6334Z"> </path> </svg> <span class="usa-sr-only">NCBI on GitHub</span> </a> <a href="https://ncbiinsights.ncbi.nlm.nih.gov/" class="ncbi-footer__social-icon ncbi-footer__social-icon--gray" target="_blank" rel="noreferrer noopener"> <svg width="26" height="26" viewBox="0 0 27 27" fill="none" xmlns="http://www.w3.org/2000/svg" focusable="false" aria-hidden="true"> <path d="M23.7778 26.4574C23.1354 26.3913 22.0856 26.8024 21.636 26.3087C21.212 25.8444 21.4359 24.8111 21.324 24.0347C19.9933 14.8323 14.8727 8.80132 6.09057 5.85008C4.37689 5.28406 2.58381 4.99533 0.779072 4.99481C0.202773 4.99481 -0.0229751 4.83146 0.00455514 4.21479C0.0660406 3.08627 0.0660406 1.95525 0.00455514 0.826734C-0.0413285 0.0815827 0.259669 -0.0193618 0.896534 0.00266238C6.96236 0.222904 12.3693 2.24179 16.9889 6.16209C22.9794 11.2478 26.1271 17.7688 26.4372 25.648C26.4629 26.294 26.3179 26.5271 25.6609 26.4684C25.0827 26.417 24.4991 26.4574 23.7778 26.4574Z"> </path> <path d="M14.8265 26.441C14.0924 26.441 13.2371 26.6795 12.6626 26.3786C12.0092 26.0372 12.3781 25.0644 12.246 24.378C11.1154 18.5324 6.6849 14.5497 0.74755 14.1001C0.217135 14.0615 -0.0104482 13.9422 0.0134113 13.3659C0.0519536 12.1454 0.0482829 10.9213 0.0134113 9.69524C-0.00127145 9.14464 0.196946 9.03268 0.703502 9.04736C9.21217 9.27128 16.5994 16.2511 17.2804 24.7231C17.418 26.4446 17.418 26.4446 15.6579 26.4446H14.832L14.8265 26.441Z"> </path> <path d="M3.58928 26.4555C2.64447 26.4618 1.73584 26.0925 1.06329 25.4289C0.39073 24.7653 0.00933763 23.8617 0.0030097 22.9169C-0.00331824 21.9721 0.365937 21.0635 1.02954 20.3909C1.69315 19.7184 2.59675 19.337 3.54156 19.3306C4.48637 19.3243 5.39499 19.6936 6.06755 20.3572C6.7401 21.0208 7.1215 21.9244 7.12782 22.8692C7.13415 23.814 6.7649 24.7226 6.10129 25.3952C5.43768 26.0677 4.53409 26.4491 3.58928 26.4555Z"> </path> </svg> <span class="usa-sr-only">NCBI RSS feed</span> </a> </div> </div> <div data-testid="gridContainer" class="grid-container ncbi-footer__container"> <div class="grid-row ncbi-footer__main-content-container" data-testid="grid"> <div class="ncbi-footer__column"> <p class="ncbi-footer__circled-icons-heading"> Connect with NLM </p> <div class="ncbi-footer__circled-icons-list"> <a href=https://twitter.com/nlm_nih class="ncbi-footer__social-icon ncbi-footer__social-icon--circled" target="_blank" rel="noreferrer noopener"> <svg width="32" height="32" viewBox="0 0 40 40" fill="none" xmlns="http://www.w3.org/2000/svg" focusable="false" aria-hidden="true"> <path d="m6.067 8 10.81 13.9L6 33.2h4.2l8.4-9.1 7.068 9.1H34L22.8 18.5 31.9 8h-3.5l-7.7 8.4L14.401 8H6.067Zm3.6 1.734h3.266l16.8 21.732H26.57L9.668 9.734Z"> </path> </svg> <span class="usa-sr-only">NLM on X (formerly known as Twitter)</span> </a> <a href=https://www.facebook.com/nationallibraryofmedicine class="ncbi-footer__social-icon ncbi-footer__social-icon--circled" target="_blank" rel="noreferrer noopener"> <svg width="13" height="24" viewBox="0 0 13 24" fill="none" xmlns="http://www.w3.org/2000/svg" focusable="false" aria-hidden="true"> <path d="M4.11371 23.1369C4.11371 23.082 4.11371 23.0294 4.11371 22.9745V12.9411H0.817305C0.6709 12.9411 0.670898 12.9411 0.670898 12.8016C0.670898 11.564 0.670898 10.3287 0.670898 9.09341C0.670898 8.97903 0.705213 8.95158 0.815017 8.95158C1.8673 8.95158 2.91959 8.95158 3.97417 8.95158H4.12057V8.83263C4.12057 7.8055 4.12057 6.7738 4.12057 5.74897C4.1264 4.92595 4.31387 4.11437 4.66959 3.37217C5.12916 2.38246 5.94651 1.60353 6.95717 1.1921C7.64827 0.905008 8.3913 0.764035 9.13953 0.778051C10.0019 0.791777 10.8644 0.830666 11.7268 0.860404C11.8869 0.860404 12.047 0.894717 12.2072 0.90158C12.2964 0.90158 12.3261 0.940469 12.3261 1.02968C12.3261 1.5421 12.3261 2.05452 12.3261 2.56465C12.3261 3.16857 12.3261 3.7725 12.3261 4.37642C12.3261 4.48165 12.2964 4.51367 12.1912 4.51138C11.5369 4.51138 10.8804 4.51138 10.2261 4.51138C9.92772 4.51814 9.63058 4.5526 9.33855 4.61433C9.08125 4.6617 8.84537 4.78881 8.66431 4.97766C8.48326 5.16652 8.3662 5.40755 8.32972 5.66661C8.28476 5.89271 8.26027 6.1224 8.25652 6.35289C8.25652 7.19014 8.25652 8.02969 8.25652 8.86923C8.25652 8.89439 8.25652 8.91955 8.25652 8.95615H12.0219C12.1797 8.95615 12.182 8.95616 12.1614 9.10714C12.0768 9.76596 11.9876 10.4248 11.9029 11.0813C11.8312 11.6319 11.7626 12.1824 11.697 12.733C11.6719 12.9434 11.6787 12.9434 11.4683 12.9434H8.26338V22.899C8.26338 22.979 8.26338 23.0591 8.26338 23.1392L4.11371 23.1369Z"> </path> </svg> <span class="usa-sr-only">NLM on Facebook</span> </a> <a href=https://www.youtube.com/user/NLMNIH class="ncbi-footer__social-icon ncbi-footer__social-icon--circled" target="_blank" rel="noreferrer noopener"> <svg width="21" height="15" viewBox="0 0 21 15" fill="none" xmlns="http://www.w3.org/2000/svg" focusable="false" aria-hidden="true"> <path d="M19.2561 1.47914C18.9016 1.15888 18.5699 0.957569 17.2271 0.834039C15.5503 0.678484 13.2787 0.655608 11.563 0.65332H9.43556C7.71987 0.65332 5.4483 0.678484 3.77151 0.834039C2.43098 0.957569 2.097 1.15888 1.74242 1.47914C0.813665 2.32097 0.619221 4.62685 0.598633 6.89384C0.598633 7.31781 0.598633 7.74101 0.598633 8.16345C0.626084 10.4121 0.827391 12.686 1.74242 13.521C2.097 13.8412 2.4287 14.0425 3.77151 14.1661C5.4483 14.3216 7.71987 14.3445 9.43556 14.3468H11.563C13.2787 14.3468 15.5503 14.3216 17.2271 14.1661C18.5676 14.0425 18.9016 13.8412 19.2561 13.521C20.1712 12.6929 20.3725 10.451 20.3999 8.22064C20.3999 7.74025 20.3999 7.25986 20.3999 6.77946C20.3725 4.54907 20.1689 2.30724 19.2561 1.47914ZM8.55942 10.5311V4.65201L13.5601 7.50005L8.55942 10.5311Z" fill="white" /> </svg> <span class="usa-sr-only">NLM on YouTube</span> </a> </div> </div> <address class="ncbi-footer__address ncbi-footer__column"> <p> <a class="usa-link usa-link--external" href="https://www.google.com/maps/place/8600+Rockville+Pike,+Bethesda,+MD+20894/%4038.9959508, -77.101021,17z/data%3D!3m1!4b1!4m5!3m4!1s0x89b7c95e25765ddb%3A0x19156f88b27635b8!8m2!3d38.9959508! 4d-77.0988323" rel="noopener noreferrer" target="_blank">National Library of Medicine <br/> 8600 Rockville Pike<br/> Bethesda, MD 20894</a> </p> </address> <ul class="usa-list usa-list--unstyled ncbi-footer__vertical-list ncbi-footer__column"> <li class="ncbi-footer__vertical-list-item"> <a href="https://www.nlm.nih.gov/web_policies.html" class="usa-link usa-link--alt ncbi-footer__link" > Web Policies </a> </li> <li class="ncbi-footer__vertical-list-item"> <a href="https://www.nih.gov/institutes-nih/nih-office-director/office-communications-public-liaison/freedom-information-act-office" class="usa-link usa-link--alt ncbi-footer__link" > FOIA </a> </li> <li class="ncbi-footer__vertical-list-item"> <a href="https://www.hhs.gov/vulnerability-disclosure-policy/index.html" class="usa-link usa-link--external usa-link--alt ncbi-footer__link" rel="noreferrer noopener" target='_blank' > HHS Vulnerability Disclosure </a> </li> </ul> <ul class="usa-list usa-list--unstyled ncbi-footer__vertical-list ncbi-footer__column"> <li class="ncbi-footer__vertical-list-item"> <a href="https://support.nlm.nih.gov/" class="usa-link usa-link--alt ncbi-footer__link" > Help </a> </li> <li class="ncbi-footer__vertical-list-item"> <a href="https://www.nlm.nih.gov/accessibility.html" class="usa-link usa-link--alt ncbi-footer__link" > Accessibility </a> </li> <li class="ncbi-footer__vertical-list-item"> <a href="https://www.nlm.nih.gov/careers/careers.html" class="usa-link usa-link--alt ncbi-footer__link" > Careers </a> </li> </ul> </div> <div class="grid-row grid-col-12" data-testid="grid"> <ul class="ncbi-footer__bottom-links-list"> <li class="ncbi-footer__bottom-list-item"> <a href="https://www.nlm.nih.gov/" class="usa-link usa-link--alt ncbi-footer__link" > NLM </a> </li> <li class="ncbi-footer__bottom-list-item"> <a href="https://www.nih.gov/" class="usa-link usa-link--alt ncbi-footer__link" > NIH </a> </li> <li class="ncbi-footer__bottom-list-item"> <a href="https://www.hhs.gov/" class="usa-link usa-link--external usa-link--alt ncbi-footer__link" rel="noreferrer noopener" target='_blank' > HHS </a> </li> <li class="ncbi-footer__bottom-list-item"> <a href="https://www.usa.gov/" class="usa-link usa-link--external usa-link--alt ncbi-footer__link" rel="noreferrer noopener" target='_blank' > USA.gov </a> </li> </ul> </div> </div> </footer> <script type="text/javascript" src="https://cdn.ncbi.nlm.nih.gov/core/pinger/pinger.js"> </script> <button class="back-to-top" data-ga-category="pagination" data-ga-action="back_to_top"> <label>Back to Top</label> <svg class="usa-icon order-0" aria-hidden="true" focusable="false" role="img"> <use xlink:href="/static/img/sprite.svg#arrow_upward"></use> </svg> </button> <script src="https://code.jquery.com/jquery-3.5.0.min.js" integrity="sha256-xNzN2a4ltkB44Mc/Jz3pT4iU1cmeR0FkXs4pru/JxaQ=" crossorigin="anonymous"> </script> <script type="text/javascript">var exports = {};</script> <script src="/static/CACHE/js/output.13b077bc3ffd.js"></script> <script type="application/javascript"> window.ncbi = window.ncbi || {}; window.ncbi.pmc = window.ncbi.pmc || {}; window.ncbi.pmc.options = { logLevel: 'INFO', staticEndpoint: '/static/', citeCookieName: 'pmc-cf', }; </script> <script type="module" crossorigin="" src="/static/assets/base-9bea7450.js"></script> <script type="module" crossorigin="" src="/static/assets/article-722d91a2.js"></script> </body> </html>

CINXE.COM

Spatially limited pathogen pollution in an invasive tick and host system - PMC