{"id":25,"date":"2019-07-25T15:25:59","date_gmt":"2019-07-25T15:25:59","guid":{"rendered":"https:\/\/www.davidpfenniglab.com\/?page_id=25"},"modified":"2021-02-01T17:55:05","modified_gmt":"2021-02-01T17:55:05","slug":"spadefoots","status":"publish","type":"page","link":"https:\/\/www.davidpfenniglab.com\/spadefoots\/","title":{"rendered":"Spadefoots!"},"content":{"rendered":"\t\t<div data-elementor-type=\"wp-page\" data-elementor-id=\"25\" class=\"elementor elementor-25\" data-elementor-post-type=\"page\">\n\t\t\t\t\t\t<section class=\"elementor-section elementor-top-section elementor-element elementor-element-ee1a795 elementor-section-stretched elementor-section-full_width elementor-section-height-default elementor-section-height-default\" data-id=\"ee1a795\" data-element_type=\"section\" data-settings=\"{"stretch_section":"section-stretched","background_background":"classic"}\">\n\t\t\t\t\t\t<div class=\"elementor-container elementor-column-gap-default\">\n\t\t\t\t\t<div class=\"elementor-column elementor-col-100 elementor-top-column elementor-element elementor-element-4a234d1\" data-id=\"4a234d1\" data-element_type=\"column\" data-settings=\"{"background_background":"classic"}\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t<div class=\"elementor-background-overlay\"><\/div>\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-d0f97cc elementor-widget elementor-widget-heading\" data-id=\"d0f97cc\" data-element_type=\"widget\" data-widget_type=\"heading.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t<h2 class=\"elementor-heading-title elementor-size-default\">Spadefoots!<\/h2>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-fee0e86 elementor-widget elementor-widget-text-editor\" data-id=\"fee0e86\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><em>An overview of their ecology, evolution, development, genetics, and behavior<\/em><\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/section>\n\t\t\t\t<section class=\"elementor-section elementor-top-section elementor-element elementor-element-6bafc27 elementor-section-boxed elementor-section-height-default elementor-section-height-default elementor-invisible\" data-id=\"6bafc27\" data-element_type=\"section\" data-settings=\"{"background_background":"classic","animation":"fadeIn"}\">\n\t\t\t\t\t\t<div class=\"elementor-container elementor-column-gap-default\">\n\t\t\t\t\t<div class=\"elementor-column elementor-col-100 elementor-top-column elementor-element elementor-element-4469b9b\" data-id=\"4469b9b\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<section class=\"elementor-section elementor-inner-section elementor-element elementor-element-2f0e0f0 elementor-section-boxed elementor-section-height-default elementor-section-height-default\" data-id=\"2f0e0f0\" data-element_type=\"section\">\n\t\t\t\t\t\t<div class=\"elementor-container elementor-column-gap-default\">\n\t\t\t\t\t<div class=\"elementor-column elementor-col-50 elementor-inner-column elementor-element elementor-element-0b13aae\" data-id=\"0b13aae\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-260fe19 elementor-widget elementor-widget-heading\" data-id=\"260fe19\" data-element_type=\"widget\" data-widget_type=\"heading.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t<h2 class=\"elementor-heading-title elementor-size-default\"><h2 class=\"elementor-heading-title elementor-size-default elementor-inline-editing pen\" data-elementor-setting-key=\"title\" data-pen-placeholder=\"Type Here...\">What is a spadefoot?<\/h2><\/h2>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-13c15c3 elementor-widget elementor-widget-text-editor\" data-id=\"13c15c3\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p>The first thing you need to know about spadefoots toads (hereafter, \u201cspadefoots\u201d) is that they\u2019re not actually toads \u2013\u2013 they\u2019re frogs. True toads are restricted to the genus <em>Bufo<\/em> (<em>Anaxyrus<\/em>). \u00a0Spadefoots get their name from the keratinized knob (or \u201cspade\u201d) on their hind feet that they use for digging (<strong>Figure 1<\/strong>).<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t<div class=\"elementor-column elementor-col-50 elementor-inner-column elementor-element elementor-element-49c9ef4\" data-id=\"49c9ef4\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-4b2b8fc elementor-widget elementor-widget-image\" data-id=\"4b2b8fc\" data-element_type=\"widget\" data-widget_type=\"image.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t\t\t\t\t\t\t<img fetchpriority=\"high\" decoding=\"async\" width=\"800\" height=\"534\" src=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-1.-Spadefoot-calling-and-burrowing-1024x683.jpg\" class=\"attachment-large size-large wp-image-730\" alt=\"\" srcset=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-1.-Spadefoot-calling-and-burrowing-1024x683.jpg 1024w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-1.-Spadefoot-calling-and-burrowing-300x200.jpg 300w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-1.-Spadefoot-calling-and-burrowing-768x512.jpg 768w\" sizes=\"(max-width: 800px) 100vw, 800px\" \/>\t\t\t\t\t\t\t\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-6bfc18e elementor-widget elementor-widget-text-editor\" data-id=\"6bfc18e\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><strong>Figure 1<\/strong>. Spadefoots, such as this Mexican spadefoot (<em>Spea multiplicata<\/em>), are equipped with numerous adaptations for living in dry environments, including their keratinized \u201cspade\u201d (arrow), which enables them to burrow underground to escape desiccation (inset). This male is calling to attract a mate. Photo taken near Portal, Arizona.<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/section>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/section>\n\t\t\t\t<section class=\"elementor-section elementor-top-section elementor-element elementor-element-41c7825 elementor-section-stretched elementor-section-full_width elementor-section-height-default elementor-section-height-default elementor-invisible\" data-id=\"41c7825\" data-element_type=\"section\" data-settings=\"{"background_background":"classic","animation":"fadeIn","stretch_section":"section-stretched"}\">\n\t\t\t\t\t\t<div class=\"elementor-container elementor-column-gap-default\">\n\t\t\t\t\t<div class=\"elementor-column elementor-col-100 elementor-top-column elementor-element elementor-element-54d7124\" data-id=\"54d7124\" data-element_type=\"column\" data-settings=\"{"background_background":"classic"}\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t<div class=\"elementor-background-overlay\"><\/div>\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-40ff80c elementor-widget elementor-widget-heading\" data-id=\"40ff80c\" data-element_type=\"widget\" data-widget_type=\"heading.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t<h2 class=\"elementor-heading-title elementor-size-default\">Where do we find spadefoots?<\/h2>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-b8a0208 elementor-widget elementor-widget-text-editor\" data-id=\"b8a0208\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p style=\"text-align: left;\"><b>Spadefoots are found in both the Old World and the New World.<\/b> The European spadefoots (family Pelobatidae) consist of one genus\u00a0(<em>Pelobates<\/em>) containing four species that are native to Europe, northwestern Africa, and western Asia.\u00a0The American spadefoots (family Scaphiopodidae) consist of two genera (<em>Scaphiopus <\/em>[3 species] and <em>Spea <\/em>[4 species]), which occur throughout North America (<strong>Figure 2<\/strong>).<\/p><p style=\"text-align: left;\">Two species\u2013\u2013<em>Scaphiopus holbrookii<\/em> and <em>Sc. hurterii<\/em>\u2013\u2013are native to eastern and central North America (<strong>Figures 2, 3<\/strong>), whereas the remaining five species\u2013\u2013<em>Sc. couchii, Spea multiplicata, Sp. bombifrons, Sp. hammondii, <\/em>and <em>Sp. intermontana<\/em>\u2013\u2013occur in arid western North America (<strong>Figures 2, 4<\/strong>). Indeed, spadefoots are unusual among amphibians in that they\u2019ve found a way to live in deserts.<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-6bda5a7 elementor-widget elementor-widget-image\" data-id=\"6bda5a7\" data-element_type=\"widget\" data-widget_type=\"image.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t\t\t\t\t\t\t<img decoding=\"async\" width=\"800\" height=\"598\" src=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Screen-Shot-2019-10-21-at-9.59.37-AM-1024x766.png\" class=\"attachment-large size-large wp-image-744\" alt=\"Figure 2: Spadefoots\" srcset=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Screen-Shot-2019-10-21-at-9.59.37-AM-1024x766.png 1024w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Screen-Shot-2019-10-21-at-9.59.37-AM-300x224.png 300w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Screen-Shot-2019-10-21-at-9.59.37-AM-768x575.png 768w\" sizes=\"(max-width: 800px) 100vw, 800px\" \/>\t\t\t\t\t\t\t\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-52a7ed4 elementor-widget elementor-widget-text-editor\" data-id=\"52a7ed4\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p style=\"text-align: center;\"><strong>Figure 2. <\/strong>Diversity and distributions of American spadefoots.<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/section>\n\t\t\t\t<section class=\"elementor-section elementor-top-section elementor-element elementor-element-989eb7f elementor-section-boxed elementor-section-height-default elementor-section-height-default elementor-invisible\" data-id=\"989eb7f\" data-element_type=\"section\" data-settings=\"{"background_background":"classic","animation":"fadeIn"}\">\n\t\t\t\t\t\t<div class=\"elementor-container elementor-column-gap-default\">\n\t\t\t\t\t<div class=\"elementor-column elementor-col-100 elementor-top-column elementor-element elementor-element-b18a21c\" data-id=\"b18a21c\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-1f4a8a7 elementor-widget elementor-widget-heading\" data-id=\"1f4a8a7\" data-element_type=\"widget\" data-widget_type=\"heading.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t<h2 class=\"elementor-heading-title elementor-size-default\">What does it look like where spadefoots live?<\/h2>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<section class=\"elementor-section elementor-inner-section elementor-element elementor-element-d5e7798 elementor-section-boxed elementor-section-height-default elementor-section-height-default\" data-id=\"d5e7798\" data-element_type=\"section\">\n\t\t\t\t\t\t<div class=\"elementor-container elementor-column-gap-default\">\n\t\t\t\t\t<div class=\"elementor-column elementor-col-50 elementor-inner-column elementor-element elementor-element-e3672c7\" data-id=\"e3672c7\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-d07dd02 elementor-widget elementor-widget-image\" data-id=\"d07dd02\" data-element_type=\"widget\" data-widget_type=\"image.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t\t\t\t\t\t\t<img decoding=\"async\" width=\"800\" height=\"602\" src=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-3.-NC-Sandhills-Pond-with-tads-1024x771.png\" class=\"attachment-large size-large wp-image-773\" alt=\"\" srcset=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-3.-NC-Sandhills-Pond-with-tads-1024x771.png 1024w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-3.-NC-Sandhills-Pond-with-tads-300x226.png 300w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-3.-NC-Sandhills-Pond-with-tads-768x578.png 768w\" sizes=\"(max-width: 800px) 100vw, 800px\" \/>\t\t\t\t\t\t\t\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-37713d2 elementor-widget elementor-widget-text-editor\" data-id=\"37713d2\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><strong>Figure 3<\/strong>. An ephemeral pond near Hoffman, North Carolina, with tadpoles (inset) of Eastern spadefoots (<em>Scaphiopus holbrookii<\/em>).<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t<div class=\"elementor-column elementor-col-50 elementor-inner-column elementor-element elementor-element-7708dd5\" data-id=\"7708dd5\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-d27cdb6 elementor-widget elementor-widget-image\" data-id=\"d27cdb6\" data-element_type=\"widget\" data-widget_type=\"image.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t\t\t\t\t\t\t<img loading=\"lazy\" decoding=\"async\" width=\"800\" height=\"600\" src=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-4.-Arizona-Pond-1024x768.jpeg\" class=\"attachment-large size-large wp-image-732\" alt=\"\" srcset=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-4.-Arizona-Pond-1024x768.jpeg 1024w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-4.-Arizona-Pond-300x225.jpeg 300w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-4.-Arizona-Pond-768x576.jpeg 768w\" sizes=\"(max-width: 800px) 100vw, 800px\" \/>\t\t\t\t\t\t\t\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-d14ecf7 elementor-widget elementor-widget-text-editor\" data-id=\"d14ecf7\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><strong>Figure 4<\/strong>. A spadefoot breeding pond in southeastern Arizona, which filled hours before this photo was taken (the storm looms over the Chiricahua Mountains).\u00a0 Spadefoots emerge from their underground burrows for only a few weeks each year to feed and breed in these temporary, rain-filled ponds. <em>Spea multiplicata <\/em>and <em>Sc. couchii <\/em>breed in this pond.<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/section>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/section>\n\t\t\t\t<section class=\"elementor-section elementor-top-section elementor-element elementor-element-d2cf955 elementor-section-stretched elementor-section-full_width elementor-section-height-default elementor-section-height-default elementor-invisible\" data-id=\"d2cf955\" data-element_type=\"section\" data-settings=\"{"background_background":"classic","animation":"fadeIn","stretch_section":"section-stretched"}\">\n\t\t\t\t\t\t<div class=\"elementor-container elementor-column-gap-default\">\n\t\t\t\t\t<div class=\"elementor-column elementor-col-100 elementor-top-column elementor-element elementor-element-60d2e13\" data-id=\"60d2e13\" data-element_type=\"column\" data-settings=\"{"background_background":"classic"}\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t<div class=\"elementor-background-overlay\"><\/div>\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-871dd52 elementor-widget elementor-widget-heading\" data-id=\"871dd52\" data-element_type=\"widget\" data-widget_type=\"heading.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t<h2 class=\"elementor-heading-title elementor-size-default\">How can spadefoots live in such extreme environments?<\/h2>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-8a57c68 elementor-widget elementor-widget-text-editor\" data-id=\"8a57c68\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><strong>Spadefoots <\/strong><strong>have evolved numerous adaptations <\/strong><strong>to cope with arid <\/strong><strong>(and often unpredictable) environments.<\/strong> These adaptations include prolonged dormancy, rapid growth and development, phenotypic plasticity, and adaptive mating between species (hybridization). <\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-6e2ac3b elementor-widget-divider--view-line elementor-widget elementor-widget-divider\" data-id=\"6e2ac3b\" data-element_type=\"widget\" data-widget_type=\"divider.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t<div class=\"elementor-divider\">\n\t\t\t<span class=\"elementor-divider-separator\">\n\t\t\t\t\t\t<\/span>\n\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<section class=\"elementor-section elementor-inner-section elementor-element elementor-element-643bf97 elementor-section-full_width elementor-section-height-default elementor-section-height-default elementor-invisible\" data-id=\"643bf97\" data-element_type=\"section\" data-settings=\"{"animation":"fadeIn"}\">\n\t\t\t\t\t\t<div class=\"elementor-container elementor-column-gap-wider\">\n\t\t\t\t\t<div class=\"elementor-column elementor-col-50 elementor-inner-column elementor-element elementor-element-aeaaae9\" data-id=\"aeaaae9\" data-element_type=\"column\" data-settings=\"{"background_background":"classic"}\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t<div class=\"elementor-background-overlay\"><\/div>\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-cb1be36 elementor-widget elementor-widget-heading\" data-id=\"cb1be36\" data-element_type=\"widget\" data-widget_type=\"heading.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t<h2 class=\"elementor-heading-title elementor-size-default\">Adaptation No. 1<\/h2>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-f87b608 elementor-widget elementor-widget-heading\" data-id=\"f87b608\" data-element_type=\"widget\" data-widget_type=\"heading.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t<h2 class=\"elementor-heading-title elementor-size-default\">Prolonged Dormancy<\/h2>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-3b7ddf3 elementor-widget elementor-widget-text-editor\" data-id=\"3b7ddf3\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p>Spadefoots are behaviorally and physiologically equipped to survive in arid environments. Adults escape dry conditions by burrowing underground using their spade (see <strong>Figure 1<\/strong>). Some species burrow over a half a meter deep!<\/p><p>Once underground, spadefoots can remain dormant for a year or longer. During this prolonged dormancy, they reduce their metabolism and conserve water by storing waste as urea and by encasing themselves in multiple protective layers of unshed skin.<\/p><p>They emerge from these burrows following warm rains (<strong>Figures 5, 6<\/strong>), which they may detect by hearing the low-frequency sound (< 100 Hz) that characterizes heavy rain striking the surface of the ground above them.<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t<div class=\"elementor-column elementor-col-50 elementor-inner-column elementor-element elementor-element-cbe82ad\" data-id=\"cbe82ad\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-6631361 elementor-widget elementor-widget-image\" data-id=\"6631361\" data-element_type=\"widget\" data-widget_type=\"image.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t\t\t\t\t\t\t<img loading=\"lazy\" decoding=\"async\" width=\"800\" height=\"600\" src=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-5.-Spadefoot-arid-habitat-with-storm-1024x768.jpeg\" class=\"attachment-large size-large wp-image-800\" alt=\"\" srcset=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-5.-Spadefoot-arid-habitat-with-storm-1024x768.jpeg 1024w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-5.-Spadefoot-arid-habitat-with-storm-300x225.jpeg 300w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-5.-Spadefoot-arid-habitat-with-storm-768x576.jpeg 768w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-5.-Spadefoot-arid-habitat-with-storm.jpeg 1280w\" sizes=\"(max-width: 800px) 100vw, 800px\" \/>\t\t\t\t\t\t\t\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-11455b3 elementor-widget elementor-widget-text-editor\" data-id=\"11455b3\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><strong>Figure 5.<\/strong> Typical habitat of <em>Sc. couchii<\/em>, Sp<em>. bombifrons<\/em>, <em>Sp. multiplicata <\/em>in southeastern Arizona. Spadefoots live underground for much of the year in a prolonged dormancy, emerging for only a few weeks following warm rains (such as this storm) to breed and feed.<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-a92826a elementor-widget elementor-widget-image\" data-id=\"a92826a\" data-element_type=\"widget\" data-widget_type=\"image.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t\t\t\t\t\t\t<img loading=\"lazy\" decoding=\"async\" width=\"800\" height=\"577\" src=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-6.-Eastern-spadefoot-emerging-from-burrow-1024x738.jpeg\" class=\"attachment-large size-large wp-image-801\" alt=\"\" srcset=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-6.-Eastern-spadefoot-emerging-from-burrow-1024x738.jpeg 1024w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-6.-Eastern-spadefoot-emerging-from-burrow-300x216.jpeg 300w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-6.-Eastern-spadefoot-emerging-from-burrow-768x553.jpeg 768w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-6.-Eastern-spadefoot-emerging-from-burrow.jpeg 1280w\" sizes=\"(max-width: 800px) 100vw, 800px\" \/>\t\t\t\t\t\t\t\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-0dc0cc3 elementor-widget elementor-widget-text-editor\" data-id=\"0dc0cc3\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><strong>Figure 6<\/strong>. <em>Scaphiopus holbrookii <\/em>emerging from its burrow following a rainstorm. <\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/section>\n\t\t\t\t<div class=\"elementor-element elementor-element-8c76844 elementor-widget-divider--view-line elementor-widget elementor-widget-divider\" data-id=\"8c76844\" data-element_type=\"widget\" data-widget_type=\"divider.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t<div class=\"elementor-divider\">\n\t\t\t<span class=\"elementor-divider-separator\">\n\t\t\t\t\t\t<\/span>\n\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<section class=\"elementor-section elementor-inner-section elementor-element elementor-element-43133be elementor-section-full_width elementor-section-height-default elementor-section-height-default elementor-invisible\" data-id=\"43133be\" data-element_type=\"section\" data-settings=\"{"animation":"fadeIn"}\">\n\t\t\t\t\t\t<div class=\"elementor-container elementor-column-gap-wider\">\n\t\t\t\t\t<div class=\"elementor-column elementor-col-50 elementor-inner-column elementor-element elementor-element-d0e57e6\" data-id=\"d0e57e6\" data-element_type=\"column\" data-settings=\"{"background_background":"classic"}\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t<div class=\"elementor-background-overlay\"><\/div>\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-848e0ee elementor-widget elementor-widget-heading\" data-id=\"848e0ee\" data-element_type=\"widget\" data-widget_type=\"heading.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t<h2 class=\"elementor-heading-title elementor-size-default\">Adaptation No. 2<\/h2>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-da4aea6 elementor-widget elementor-widget-heading\" data-id=\"da4aea6\" data-element_type=\"widget\" data-widget_type=\"heading.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t<h2 class=\"elementor-heading-title elementor-size-default\">Rapid Growth and Development\n<\/h2>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-a514ab7 elementor-widget elementor-widget-text-editor\" data-id=\"a514ab7\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p>Regardless of whether or not they live in the desert, spadefoots breed in temporary, rain-filled ponds (see <strong>Figures 3, 4<\/strong>).\u00a0 Although most frogs and toads could not reproduce in such highly ephemeral environments, spadefoot tadpoles can survive in them by developing rapidly\u2013\u2013in some cases metamorphosing in eight days post-hatching (<strong>Figure 7<\/strong>).<\/p><p>Furthermore, spadefoot tadpoles develop lungs relatively early in life.\u00a0 Consequently, if their pond dries, they can remain alive on dry land for days (<strong>Figure 8<\/strong>), thereby increasing their chances of surviving until another rainstorm refills their pond.<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t<div class=\"elementor-column elementor-col-50 elementor-inner-column elementor-element elementor-element-34e6050\" data-id=\"34e6050\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-b04f989 elementor-widget elementor-widget-image\" data-id=\"b04f989\" data-element_type=\"widget\" data-widget_type=\"image.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t\t\t\t\t\t\t<img loading=\"lazy\" decoding=\"async\" width=\"800\" height=\"557\" src=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-5.-Metamorph-1024x713.jpeg\" class=\"attachment-large size-large wp-image-733\" alt=\"\" srcset=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-5.-Metamorph-1024x713.jpeg 1024w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-5.-Metamorph-300x209.jpeg 300w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-5.-Metamorph-768x535.jpeg 768w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-5.-Metamorph.jpeg 1895w\" sizes=\"(max-width: 800px) 100vw, 800px\" \/>\t\t\t\t\t\t\t\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-69acadf elementor-widget elementor-widget-text-editor\" data-id=\"69acadf\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><strong>Figure 7<\/strong>. Spadefoot tadpoles exhibit rapid, and adaptively flexible, larval development.\u00a0 Here, a <em>Spea multiplicata<\/em> has developed its limbs and thereby escapes its rapidly drying pond (photo near Portal, Arizona).<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-719c695 elementor-widget elementor-widget-image\" data-id=\"719c695\" data-element_type=\"widget\" data-widget_type=\"image.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t\t\t\t\t\t\t<img loading=\"lazy\" decoding=\"async\" width=\"800\" height=\"600\" src=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-6.-Tiger-beetles-eating-drying-tads-1024x768.jpeg\" class=\"attachment-large size-large wp-image-736\" alt=\"\" srcset=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-6.-Tiger-beetles-eating-drying-tads-1024x768.jpeg 1024w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-6.-Tiger-beetles-eating-drying-tads-300x225.jpeg 300w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-6.-Tiger-beetles-eating-drying-tads-768x576.jpeg 768w\" sizes=\"(max-width: 800px) 100vw, 800px\" \/>\t\t\t\t\t\t\t\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-6d0c77a elementor-widget elementor-widget-text-editor\" data-id=\"6d0c77a\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><strong>Figure 8<\/strong>. Because they develop lungs early, <em>Spea tadpoles<\/em> can survive on land as tadpoles for days. These <em>Spea multiplicata<\/em> tadpoles (encased in mud) are still alive. However, they are vulnerable to predators, such as these tiger beetles.<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/section>\n\t\t\t\t<div class=\"elementor-element elementor-element-f130a8a elementor-widget-divider--view-line elementor-widget elementor-widget-divider\" data-id=\"f130a8a\" data-element_type=\"widget\" data-widget_type=\"divider.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t<div class=\"elementor-divider\">\n\t\t\t<span class=\"elementor-divider-separator\">\n\t\t\t\t\t\t<\/span>\n\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<section class=\"elementor-section elementor-inner-section elementor-element elementor-element-b223513 elementor-section-full_width elementor-section-height-default elementor-section-height-default elementor-invisible\" data-id=\"b223513\" data-element_type=\"section\" data-settings=\"{"animation":"fadeIn"}\">\n\t\t\t\t\t\t<div class=\"elementor-container elementor-column-gap-wider\">\n\t\t\t\t\t<div class=\"elementor-column elementor-col-50 elementor-inner-column elementor-element elementor-element-604caed\" data-id=\"604caed\" data-element_type=\"column\" data-settings=\"{"background_background":"classic"}\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t<div class=\"elementor-background-overlay\"><\/div>\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-b4cb897 elementor-widget elementor-widget-heading\" data-id=\"b4cb897\" data-element_type=\"widget\" data-widget_type=\"heading.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t<h2 class=\"elementor-heading-title elementor-size-default\">Adaptation No. 3<\/h2>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-d742075 elementor-widget elementor-widget-heading\" data-id=\"d742075\" data-element_type=\"widget\" data-widget_type=\"heading.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t<h2 class=\"elementor-heading-title elementor-size-default\">Phenotypic Plasticity\n<\/h2>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-dd2bba5 elementor-widget elementor-widget-text-editor\" data-id=\"dd2bba5\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p>Spadefoots also exhibit multiple forms of phenotypic plasticity that allows them to thrive in environments (such as deserts) where rainfall is highly variable. Specifically, in response to changing water levels in their pond, spadefoot tadpoles can either speed up or slow down their development.<\/p><p>\u00a0<\/p><p>Moreover, whereas most frog and toad tadpoles are omnivores and exhibit traits adapted for feeding on detritus and plankton, tadpoles of the genus <em>Spea<\/em> can develop into an alternative\u2013\u2013and more rapidly developing\u2013\u2013carnivore morph, which exhibits enlarged jaw muscles and mouthparts for capturing and consuming large animal prey, such as fairy shrimp and other tadpoles (<strong>Figure <\/strong><strong>9<\/strong>).<\/p><p>\u00a0<\/p><p>The carnivore morph has an additional advantage in a rapidly drying pond: by eating its competitors, it can reduce competition and further enhance growth and development (<strong>Figure <\/strong><strong>10<\/strong>). However, when engaging in cannibalism, carnivores can distinguish between kin and nonkin, preferentially cannibalizing the latter.<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t<div class=\"elementor-column elementor-col-50 elementor-inner-column elementor-element elementor-element-221bb76\" data-id=\"221bb76\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-366c0c6 elementor-widget elementor-widget-image\" data-id=\"366c0c6\" data-element_type=\"widget\" data-widget_type=\"image.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t\t\t\t\t\t\t<img loading=\"lazy\" decoding=\"async\" width=\"800\" height=\"450\" src=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-7.-Alternative-tadpole-morphs-1024x576.jpeg\" class=\"attachment-large size-large wp-image-737\" alt=\"\" srcset=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-7.-Alternative-tadpole-morphs-1024x576.jpeg 1024w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-7.-Alternative-tadpole-morphs-300x169.jpeg 300w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-7.-Alternative-tadpole-morphs-768x432.jpeg 768w\" sizes=\"(max-width: 800px) 100vw, 800px\" \/>\t\t\t\t\t\t\t\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-6ed2618 elementor-widget elementor-widget-text-editor\" data-id=\"6ed2618\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><strong>Figure 9<\/strong>. <em>Spea<\/em> produce alternative, environmentally induced, tadpole morphs: a slower developing omnivore morph (left) and a more rapidly developing carnivore morph (right), which is induced by, and specializes on, animal prey, such as fairy shrimp (center).<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-4fdca35 elementor-widget elementor-widget-image\" data-id=\"4fdca35\" data-element_type=\"widget\" data-widget_type=\"image.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t\t\t\t\t\t\t<img loading=\"lazy\" decoding=\"async\" width=\"800\" height=\"617\" src=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-8.-Cannibalism-1024x790.jpeg\" class=\"attachment-large size-large wp-image-738\" alt=\"\" srcset=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-8.-Cannibalism-1024x790.jpeg 1024w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-8.-Cannibalism-300x231.jpeg 300w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-8.-Cannibalism-768x593.jpeg 768w\" sizes=\"(max-width: 800px) 100vw, 800px\" \/>\t\t\t\t\t\t\t\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-8fe1b66 elementor-widget elementor-widget-text-editor\" data-id=\"8fe1b66\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><strong>Figure 10<\/strong>. <em>Spea<\/em> carnivore-morph tadpoles are highly cannibalistic. Here, a carnivore <em>Sp. multiplicata<\/em> eats an omnivore.<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/section>\n\t\t\t\t<div class=\"elementor-element elementor-element-0cb84ba elementor-widget-divider--view-line elementor-widget elementor-widget-divider\" data-id=\"0cb84ba\" data-element_type=\"widget\" data-widget_type=\"divider.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t<div class=\"elementor-divider\">\n\t\t\t<span class=\"elementor-divider-separator\">\n\t\t\t\t\t\t<\/span>\n\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<section class=\"elementor-section elementor-inner-section elementor-element elementor-element-cabefd1 elementor-section-full_width elementor-section-height-default elementor-section-height-default elementor-invisible\" data-id=\"cabefd1\" data-element_type=\"section\" data-settings=\"{"animation":"fadeIn"}\">\n\t\t\t\t\t\t<div class=\"elementor-container elementor-column-gap-wider\">\n\t\t\t\t\t<div class=\"elementor-column elementor-col-50 elementor-inner-column elementor-element elementor-element-6610b43\" data-id=\"6610b43\" data-element_type=\"column\" data-settings=\"{"background_background":"classic"}\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t<div class=\"elementor-background-overlay\"><\/div>\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-ee1f7d7 elementor-widget elementor-widget-heading\" data-id=\"ee1f7d7\" data-element_type=\"widget\" data-widget_type=\"heading.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t<h2 class=\"elementor-heading-title elementor-size-default\">Adaptation No. 4<\/h2>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-6566cf0 elementor-widget elementor-widget-heading\" data-id=\"6566cf0\" data-element_type=\"widget\" data-widget_type=\"heading.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t<h2 class=\"elementor-heading-title elementor-size-default\">Adaptive Mating Between Species\n\n<\/h2>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-298fda0 elementor-widget elementor-widget-text-editor\" data-id=\"298fda0\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p>Finally, as adults, at least one species of spadefoot exhibits remarkable behavioral plasticity. Such plasticity appears to have enabled this species to invade desert habitats.<\/p><p>In particular, when breeding in shallow, rapidly drying ponds, <em>Sp. bombifrons<\/em>\u00a0females often switch from mating with their own species (<strong>Figure <\/strong><strong>11<\/strong>) to preferring males of\u00a0<em>Sp. multiplicata\u00a0<\/em>(<strong>Figure <\/strong><strong>12<\/strong>). In the desert southwest, these two species often breed in the same ponds, and <em>Sp. multiplicata<\/em><em>\u00a0<\/em>tadpoles develop faster than <em>Sp. bombifrons <\/em>tadpoles.<\/p><p>Because the resulting <em>Sp. bombifrons <\/em>x <em>Sp. multiplicata<\/em><em>\u00a0<\/em>hybrid tadpoles (which are viable) develop faster than pure-<em>Sp. bombifrons<\/em>\u00a0tadpoles, they are more likely to escape a drying pond.\u00a0 As long as some of these hybrid offspring mature and ultimately reproduce successfully (as genetic evidence suggests that they do), this form of behavioral plasticity can be an adaptive\u2013\u2013albeit extreme\u2013\u2013strategy for dealing with dry environments.<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t<div class=\"elementor-column elementor-col-50 elementor-inner-column elementor-element elementor-element-4612f20\" data-id=\"4612f20\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-38ecb3c elementor-widget elementor-widget-image\" data-id=\"38ecb3c\" data-element_type=\"widget\" data-widget_type=\"image.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t\t\t\t\t\t\t<img loading=\"lazy\" decoding=\"async\" width=\"800\" height=\"560\" src=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-9.-Spea-bombifrons-pair-1024x717.jpeg\" class=\"attachment-large size-large wp-image-739\" alt=\"\" srcset=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-9.-Spea-bombifrons-pair-1024x717.jpeg 1024w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-9.-Spea-bombifrons-pair-300x210.jpeg 300w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-9.-Spea-bombifrons-pair-768x538.jpeg 768w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-9.-Spea-bombifrons-pair.jpeg 2043w\" sizes=\"(max-width: 800px) 100vw, 800px\" \/>\t\t\t\t\t\t\t\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-f841f08 elementor-widget elementor-widget-text-editor\" data-id=\"f841f08\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><strong>Figure <\/strong><strong>11<\/strong>. A mated pair (male on back of female) of Plains spadefoots,\u00a0<em>Spea bombifrons<\/em>.<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-c779b0f elementor-widget elementor-widget-image\" data-id=\"c779b0f\" data-element_type=\"widget\" data-widget_type=\"image.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t\t\t\t\t\t\t<img loading=\"lazy\" decoding=\"async\" width=\"800\" height=\"623\" src=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/11\/0-1024x798.jpg\" class=\"attachment-large size-large wp-image-816\" alt=\"\" srcset=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/11\/0-1024x798.jpg 1024w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/11\/0-300x234.jpg 300w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/11\/0-768x599.jpg 768w\" sizes=\"(max-width: 800px) 100vw, 800px\" \/>\t\t\t\t\t\t\t\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-6ae62fd elementor-widget elementor-widget-text-editor\" data-id=\"6ae62fd\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><strong>Figure 12<\/strong>. A male Mexican spadefoot, <em>Sp. multiplicata <\/em>(left)<em>, <\/em>mated with a female Plains spadefoot,\u00a0<em>Spea bombifrons (right)<\/em>.<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/section>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/section>\n\t\t\t\t<section class=\"elementor-section elementor-top-section elementor-element elementor-element-fdc23e3 elementor-section-boxed elementor-section-height-default elementor-section-height-default elementor-invisible\" data-id=\"fdc23e3\" data-element_type=\"section\" data-settings=\"{"background_background":"classic","animation":"fadeIn"}\">\n\t\t\t\t\t\t\t<div class=\"elementor-background-overlay\"><\/div>\n\t\t\t\t\t\t\t<div class=\"elementor-container elementor-column-gap-default\">\n\t\t\t\t\t<div class=\"elementor-column elementor-col-100 elementor-top-column elementor-element elementor-element-5bf5ea4\" data-id=\"5bf5ea4\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-8117b45 elementor-widget elementor-widget-heading\" data-id=\"8117b45\" data-element_type=\"widget\" data-widget_type=\"heading.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t<h2 class=\"elementor-heading-title elementor-size-default\">How do we track adaptations?<\/h2>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-bb30a34 elementor-widget elementor-widget-text-editor\" data-id=\"bb30a34\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><strong>To help begin to identify the genetic bases of spadefoots\u2019 unique adaptations, we recently sequenced the genome of<\/strong><strong>\u00a0<\/strong><em><strong>Spea multiplicata<\/strong><\/em><strong>.<\/strong>\u00a0<\/p><p>One discovery that we\u2019ve already made is that <em>Sp. multiplicata<\/em><em>\u00a0<\/em>possesses one of the smallest amphibian genomes known (<strong>Figure 13A<\/strong>).\u00a0 Moreover, compared to the genomes of other sequenced frog and toad species, the genome of <em>Sp. multiplicata<\/em> contains less repetitive DNA (DNA\u00a0sequences that are repeated in the genome and that do not code for protein; <strong>Figure 13B<\/strong>). Additionally, relative to these other sequenced frog and toad species, <em>Sp. multiplicata<\/em><em>\u00a0<\/em>also has smaller genes (<strong>Figure 13C<\/strong>). The fact that the different species differ in gene length, but not in transcript length (see <strong>Figure 13C<\/strong>), suggests that <em>Sp. multiplicata <\/em>has smaller genes because some non-coding sequences have been purged from their genes.<\/p><p>Such a reduction in genome and gene size makes sense. As noted above, <em>Sp. multiplicata<\/em><em>\u00a0<\/em>experiences strong selection to develop rapidly: one way to speed development is to have less DNA to replicate!<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<section class=\"elementor-section elementor-inner-section elementor-element elementor-element-503e63d elementor-section-boxed elementor-section-height-default elementor-section-height-default\" data-id=\"503e63d\" data-element_type=\"section\">\n\t\t\t\t\t\t<div class=\"elementor-container elementor-column-gap-default\">\n\t\t\t\t\t<div class=\"elementor-column elementor-col-50 elementor-inner-column elementor-element elementor-element-f812cc3\" data-id=\"f812cc3\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-523c29c elementor-widget elementor-widget-image\" data-id=\"523c29c\" data-element_type=\"widget\" data-widget_type=\"image.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t\t\t\t\t\t\t<img loading=\"lazy\" decoding=\"async\" width=\"800\" height=\"290\" src=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-12.-Genome-1024x371.png\" class=\"attachment-large size-large wp-image-802\" alt=\"\" srcset=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-12.-Genome-1024x371.png 1024w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-12.-Genome-300x109.png 300w, https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Figure-12.-Genome-768x278.png 768w\" sizes=\"(max-width: 800px) 100vw, 800px\" \/>\t\t\t\t\t\t\t\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t<div class=\"elementor-column elementor-col-50 elementor-inner-column elementor-element elementor-element-0d7b22d\" data-id=\"0d7b22d\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-b9331aa elementor-widget elementor-widget-text-editor\" data-id=\"b9331aa\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><strong>Figure 13<\/strong>. <strong>Factors contributing to differences in genome size among frogs and toads<\/strong>. <strong>A<\/strong> Estimated genome sizes of 284 species of anurans; arrow: genome size of <em>Spea<\/em>. <strong>B<\/strong> Genome (assembly) size and <strong>C<\/strong> gene and transcript lengths of <em>Sp. multiplicata <\/em>compared to three other species whose genomes have been sequenced. Figure from Seidl et al. (2019). <\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/section>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/section>\n\t\t\t\t<section class=\"elementor-section elementor-top-section elementor-element elementor-element-b728c40 elementor-section-stretched elementor-section-full_width elementor-section-height-default elementor-section-height-default elementor-invisible\" data-id=\"b728c40\" data-element_type=\"section\" data-settings=\"{"background_background":"classic","animation":"fadeIn","stretch_section":"section-stretched"}\">\n\t\t\t\t\t\t<div class=\"elementor-container elementor-column-gap-default\">\n\t\t\t\t\t<div class=\"elementor-column elementor-col-100 elementor-top-column elementor-element elementor-element-626989f\" data-id=\"626989f\" data-element_type=\"column\" data-settings=\"{"background_background":"classic"}\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t<div class=\"elementor-background-overlay\"><\/div>\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-b2bdade elementor-widget elementor-widget-heading\" data-id=\"b2bdade\" data-element_type=\"widget\" data-widget_type=\"heading.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t<h2 class=\"elementor-heading-title elementor-size-default\">Cool! How can I learn more?<\/h2>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-88e7a29 elementor-widget elementor-widget-text-editor\" data-id=\"88e7a29\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p style=\"text-align: center;\">Below are some videos and a list of scientific papers for you to learn more about these remarkable creatures.<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<section class=\"elementor-section elementor-inner-section elementor-element elementor-element-999118c elementor-section-full_width elementor-section-height-default elementor-section-height-default\" data-id=\"999118c\" data-element_type=\"section\">\n\t\t\t\t\t\t<div class=\"elementor-container elementor-column-gap-default\">\n\t\t\t\t\t<div class=\"elementor-column elementor-col-33 elementor-inner-column elementor-element elementor-element-ea49d5b\" data-id=\"ea49d5b\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-bd4a378 elementor-widget elementor-widget-video\" data-id=\"bd4a378\" data-element_type=\"widget\" data-settings=\"{"youtube_url":"https:\\\/\\\/www.youtube.com\\\/watch?v=SxxkoLfQWWA&feature=youtu.be","video_type":"youtube"}\" data-widget_type=\"video.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t<div class=\"elementor-wrapper elementor-open-inline\">\n\t\t\t<div class=\"elementor-video\"><\/div>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-b37cb56 elementor-widget elementor-widget-text-editor\" data-id=\"b37cb56\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><em>Sp. multiplicata <\/em>male calling <\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t<div class=\"elementor-column elementor-col-33 elementor-inner-column elementor-element elementor-element-b64b87a\" data-id=\"b64b87a\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-d5608f8 elementor-widget elementor-widget-video\" data-id=\"d5608f8\" data-element_type=\"widget\" data-settings=\"{"youtube_url":"https:\\\/\\\/www.youtube.com\\\/watch?v=eE6quiljb_0","video_type":"youtube","controls":"yes"}\" data-widget_type=\"video.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t<div class=\"elementor-wrapper elementor-open-inline\">\n\t\t\t<div class=\"elementor-video\"><\/div>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-137cc4d elementor-widget elementor-widget-text-editor\" data-id=\"137cc4d\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><em>Sp. multiplicata <\/em>male calling and then being chosen by a female <\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t<div class=\"elementor-column elementor-col-33 elementor-inner-column elementor-element elementor-element-79154f5\" data-id=\"79154f5\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-58bfeef elementor-widget elementor-widget-video\" data-id=\"58bfeef\" data-element_type=\"widget\" data-settings=\"{"youtube_url":"https:\\\/\\\/www.youtube.com\\\/watch?v=vsxswLkMaKg","video_type":"youtube","controls":"yes"}\" data-widget_type=\"video.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t<div class=\"elementor-wrapper elementor-open-inline\">\n\t\t\t<div class=\"elementor-video\"><\/div>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-4f9348f elementor-widget elementor-widget-text-editor\" data-id=\"4f9348f\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><em>Sp. bombifrons <\/em>male calling <\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/section>\n\t\t\t\t<div class=\"elementor-element elementor-element-532c4c4 elementor-widget-divider--view-line elementor-widget elementor-widget-divider\" data-id=\"532c4c4\" data-element_type=\"widget\" data-widget_type=\"divider.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t<div class=\"elementor-divider\">\n\t\t\t<span class=\"elementor-divider-separator\">\n\t\t\t\t\t\t<\/span>\n\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<section class=\"elementor-section elementor-inner-section elementor-element elementor-element-7fa8fc8 elementor-section-full_width elementor-section-height-default elementor-section-height-default\" data-id=\"7fa8fc8\" data-element_type=\"section\">\n\t\t\t\t\t\t<div class=\"elementor-container elementor-column-gap-default\">\n\t\t\t\t\t<div class=\"elementor-column elementor-col-33 elementor-inner-column elementor-element elementor-element-93e1dee\" data-id=\"93e1dee\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-16e63ac elementor-widget elementor-widget-video\" data-id=\"16e63ac\" data-element_type=\"widget\" data-settings=\"{"youtube_url":"https:\\\/\\\/www.youtube.com\\\/watch?v=gzja2w0W8j0&feature=youtu.be","video_type":"youtube","controls":"yes"}\" data-widget_type=\"video.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t<div class=\"elementor-wrapper elementor-open-inline\">\n\t\t\t<div class=\"elementor-video\"><\/div>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-c687f03 elementor-widget elementor-widget-text-editor\" data-id=\"c687f03\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><em>Sc. couchii <\/em>male calling <\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t<div class=\"elementor-column elementor-col-33 elementor-inner-column elementor-element elementor-element-29a22ad\" data-id=\"29a22ad\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-88716d4 elementor-widget elementor-widget-video\" data-id=\"88716d4\" data-element_type=\"widget\" data-settings=\"{"youtube_url":"https:\\\/\\\/www.youtube.com\\\/watch?v=6GoH1tmAX0A&feature=youtu.be","video_type":"youtube","controls":"yes"}\" data-widget_type=\"video.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t<div class=\"elementor-wrapper elementor-open-inline\">\n\t\t\t<div class=\"elementor-video\"><\/div>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-2216557 elementor-widget elementor-widget-text-editor\" data-id=\"2216557\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><em>Sp. multiplicata <\/em>tadpoles in the wild <\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t<div class=\"elementor-column elementor-col-33 elementor-inner-column elementor-element elementor-element-d2952ee\" data-id=\"d2952ee\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-113c714 elementor-widget elementor-widget-video\" data-id=\"113c714\" data-element_type=\"widget\" data-settings=\"{"youtube_url":"https:\\\/\\\/www.youtube.com\\\/watch?v=4EOPvZ__6Z4&feature=youtu.be","video_type":"youtube","controls":"yes"}\" data-widget_type=\"video.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t<div class=\"elementor-wrapper elementor-open-inline\">\n\t\t\t<div class=\"elementor-video\"><\/div>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-c0c1d4e elementor-widget elementor-widget-text-editor\" data-id=\"c0c1d4e\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><em>Sp. multiplicata <\/em>carnivores and omnivores from a single clutch of eggs, showing difference between morphs in behavior, even among siblings <\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/section>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/section>\n\t\t\t\t<section class=\"elementor-section elementor-top-section elementor-element elementor-element-1698a12 outreach elementor-section-boxed elementor-section-height-default elementor-section-height-default elementor-invisible\" data-id=\"1698a12\" data-element_type=\"section\" data-settings=\"{"background_background":"classic","animation":"fadeIn"}\">\n\t\t\t\t\t\t<div class=\"elementor-container elementor-column-gap-default\">\n\t\t\t\t\t<div class=\"elementor-column elementor-col-100 elementor-top-column elementor-element elementor-element-4b27ab3\" data-id=\"4b27ab3\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-81a5be8 elementor-widget elementor-widget-heading\" data-id=\"81a5be8\" data-element_type=\"widget\" data-widget_type=\"heading.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t<h2 class=\"elementor-heading-title elementor-size-default\">Here\u2019s a short (<\u00a05 min.) video on our spadefoot research:<\/h2>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-9d1b223 elementor-widget elementor-widget-video\" data-id=\"9d1b223\" data-element_type=\"widget\" data-settings=\"{"youtube_url":"https:\\\/\\\/www.youtube.com\\\/watch?v=k-AIJicehEo&feature=youtu.be","video_type":"youtube","controls":"yes"}\" data-widget_type=\"video.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t<div class=\"elementor-wrapper elementor-open-inline\">\n\t\t\t<div class=\"elementor-video\"><\/div>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-5f1161d elementor-widget elementor-widget-text-editor\" data-id=\"5f1161d\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p style=\"text-align: center;\">From the National Science Foundations\u2019s <a href=\"https:\/\/www.youtube.com\/user\/VideosatNSF\" target=\"_blank\" rel=\"noopener\">Science Nation<\/a> series.<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/section>\n\t\t\t\t<section class=\"elementor-section elementor-top-section elementor-element elementor-element-163e832 outreach elementor-section-boxed elementor-section-height-default elementor-section-height-default elementor-invisible\" data-id=\"163e832\" data-element_type=\"section\" data-settings=\"{"animation":"fadeIn"}\">\n\t\t\t\t\t\t<div class=\"elementor-container elementor-column-gap-default\">\n\t\t\t\t\t<div class=\"elementor-column elementor-col-50 elementor-top-column elementor-element elementor-element-7032219\" data-id=\"7032219\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-e79aba1 elementor-widget elementor-widget-heading\" data-id=\"e79aba1\" data-element_type=\"widget\" data-widget_type=\"heading.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t<h2 class=\"elementor-heading-title elementor-size-default\">Here are some publications on spadefoots:\n<\/h2>\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t<div class=\"elementor-element elementor-element-3f3bd22 publications elementor-widget elementor-widget-text-editor\" data-id=\"3f3bd22\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p>Bragg, A. N. 1965. Gnomes of the night: the spadefoot toads. <i><b>University of Pennsylvania Press<\/b><\/i>, Philadelphia, PA.<\/p><p><span style=\"font-weight: 400;\"><a href=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2020\/09\/Chen-and-Pfennig-2020-Science.pdf\">PDF<\/a> Chen, C. and K. S. Pfennig. 2020. Female toads engaging in adaptive hybridization prefer high-quality heterospecifics as mates. <\/span><b><i>Science<\/i><\/b><span style=\"font-weight: 400;\"> 367:1377.<\/span><\/p><p>Denver, R. J., N. Mirhadi, and M. Phillips. 1998. Adaptive plasticity in amphibian metamorphosis: response of <em>Scaphiopus hammondii<\/em> tadpoles to habitat desiccation. <i><b>Ecology<\/b><\/i> 79:1859-1872.<\/p><p>Dimmitt, M. A. & Ruibal, R. 1980. Environmental correlates of emergence in spadefoot toads (<em>Scaphiopus<\/em>). <strong><em>Journal of Herpetology<\/em><\/strong> 14: 21-29.<\/p><p>Gomez-Mestre, I. and D. R. Buchholz. 2006. Developmental plasticity mirrors differences among taxa in spadefoot toads linking plasticity and diversity. <i><b>Proceedings of the National Academy of Sciences, U.S.A.<\/b><\/i> 103:19021-19026.<\/p><p><a href=\"http:\/\/davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/9.-Kelly-et-al.-Male-sexual-signal-predicts-phenotypic-plasticity-in-offspring-implications-for-the-evolution-of-plasticity-and-local-adaptation.pdf\" target=\"_blank\" rel=\"noopener\">PDF<\/a> Kelly, P. W., D. W. Pfennig, S. de la Serna Buz\u00f3n, and K. S. Pfennig. 2019. Male sexual signals predict phenotypic plasticity in offspring: implications for the evolution of plasticity and local adaptation.<i><b> Philos. Trans. R. Soc. <\/b><\/i>B-Biol. Sci. 374:20180179.<\/p><p><a href=\"http:\/\/davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/2011_Evo_Devo.pdf\" target=\"_blank\" rel=\"noopener\">PDF<\/a> Led\u00f3n-Rettig, C. C. and D. W. Pfennig. 2011. Emerging model systems in eco-evo-devo: the environmentally responsive spadefoot toad. <i><b>Evolution and Development<\/b><\/i> 13:391-400.<\/p><p><a title=\"Publications_files\/2018_Nature_Ecology_&_Evolution.pdf\" href=\"http:\/\/davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/2018_Nature_Ecology__Evolution1.pdf\">PDF<\/a> Levis, N. A., A. Isdaner, and D. W. Pfennig. 2018. Morphological novelty emerges from pre-existing phenotypic plasticity. <i><b>Nature Ecology & Evolution<\/b><\/i> 2:1289\u20131297.<\/p><p><a title=\"Publications_files\/2017_Evolution.pdf\" href=\"http:\/\/davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/2017_Evolution.pdf\">PDF<\/a> Levis, N. A., R. A. Martin, K. A. O\u2019Donnell, and D. W. Pfennig. 2017. Intraspecific adaptive radiation: competition, ecological opportunity, and phenotypic diversification within species. <i><b>Evolution<\/b><\/i> 71:2496\u20132509.<\/p><p><a title=\"Publications_files\/2009_Am_Nat.pdf\" href=\"http:\/\/labs.bio.unc.edu\/pfennig\/LabSite\/Publications_files\/2009_Am_Nat.pdf\">PDF<\/a> Martin, R. A. and D. W. Pfennig. 2009. Disruptive selection in natural populations: the roles of ecological specialization and resource competition. <i><b>Am. Nat.<\/b><\/i> 174:268-281.<\/p><p><span style=\"font-weight: 400;\"><a href=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2020\/09\/Pfennig-and-Pfennig-2020-Copeia.pdf\">PDF<\/a> Pfennig, K. S. and Pfennig, D. W. 2020. Dead spadefoot tadpoles adaptively modify development in future generations: a novel form of nongenetic inheritance? <\/span><b><i>Copeia<\/i><\/b> <span style=\"font-weight: 400;\">108: 116-121.<\/span><\/p><p><a title=\"Publications_files\/1990_Oecologia.pdf\" href=\"http:\/\/davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/1990_Oecologia.pdf\">PDF<\/a> Pfennig, D. W. 1990. The adaptive significance of an environmentally-cued developmental switch in an anuran tadpole. <i><b>Oecologia <\/b><\/i>85:101-107.<\/p><p><a title=\"Publications_files\/1992_Evolution.pdf\" href=\"http:\/\/davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/1992_Evolution.pdf\">PDF<\/a> Pfennig, D. W. 1992. Polyphenism in spadefoot toads as a locally adjusted evolutionarily stable strategy. <i><b>Evolution <\/b><\/i>46:1408-1420.<\/p><p><a title=\"Publications_files\/2002_Evolution.pdf\" href=\"http:\/\/davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/2002_Evolution.pdf\">PDF<\/a> Pfennig, D. W. and P. J. Murphy. 2002. How fluctuating competition and phenotypic plasticity mediate species divergence. <i><b>Evolution <\/b><\/i>56:1217-1228.<\/p><p>\u00a0<\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t<div class=\"elementor-column elementor-col-50 elementor-top-column elementor-element elementor-element-17908db\" data-id=\"17908db\" data-element_type=\"column\">\n\t\t\t<div class=\"elementor-widget-wrap elementor-element-populated\">\n\t\t\t\t\t\t<div class=\"elementor-element elementor-element-d9c38bb publications elementor-widget elementor-widget-text-editor\" data-id=\"d9c38bb\" data-element_type=\"widget\" data-widget_type=\"text-editor.default\">\n\t\t\t\t<div class=\"elementor-widget-container\">\n\t\t\t\t\t\t\t<p><a href=\"http:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Pfennig-et-al.-1993_Anim_Behav.pdf\">PDF<\/a> Pfennig, D. W., H. K. Reeve, and P. W. Sherman. 1993. Kin recognition and cannibalism in spadefoot toad tadpoles. <i><b>Anim. Behav.<\/b><\/i> 46:87-94.<\/p><p><a href=\"http:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Pfennig_2000.pdf\" target=\"_blank\" rel=\"noopener\">PDF<\/a> Pfennig, K. S. 2000. Female spadefoot toads compromise on mate quality to ensure conspecific matings. <i><b>Behavioral Ecology<\/b><\/i> 11:220-227.<\/p><p><a href=\"http:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Pfennig-2007-Science.pdf\" target=\"_blank\" rel=\"noopener\">PDF<\/a> Pfennig, K. S. 2007. Facultative mate choice drives adaptive hybridization. <i><b>Science<\/b><\/i> 318:965-967.<\/p><p><a href=\"http:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Pfennig-and-Rice-2014-Reinforcement-generates-reproductive-isolation-between-neighbouring-conspecific-populations-of-spadefoot-toads.pdf\" target=\"_blank\" rel=\"noopener\">PDF<\/a> Pfennig, K. S. & Rice, A. M. 2014. Reinforcement generates reproductive isolation between neighbouring populations of spadefoot toads. Proceedings of the Royal Society B: Biological Sciences 281: 20140949.<\/p><p><a href=\"http:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Pfennig_and_Tinsley_2002.pdf\" target=\"_blank\" rel=\"noopener\">PDF<\/a> Pfennig, K. S. and R. C. Tinsley. 2002. Different mate preferences by parasitized and unparasitized females potentially reduces sexual selection.<i><b> Journal of Evolutionary Biology <\/b><\/i>15:399-406.<\/p><p><a href=\"http:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2019\/10\/Pierce-et-al.-2017-Genetic-variation-during-range-expansion-effects-of-habitat-novelty-and-hybridization.pdf\" target=\"_blank\" rel=\"noopener\">PDF<\/a> Pierce, A. A., Gutierrez, R., Rice, A. M. & Pfennig, K. S. 2017. Genetic variation during range expansion: effects of habitat novelty and hybridization. <strong><em>Proceedings of the Royal Society B: Biological Sciences<\/em><\/strong> 284: 20170007.<\/p><p>Pomeroy, L. V. 1981. Developmental polymorphism in the tadpoles of the spadefoot toad <em>Scaphiopus multiplicatus<\/em>. <i><b>University of California, Riverside, CA.<\/b><\/i><\/p><p>Ruibal, R., L. Tevis, and V. Roig. 1969. The terrestrial ecology of the spadefoot toad <em>Scaphiopus hammondii<\/em>. <i><b>Copeia<\/b><\/i> 1969:571-584.<\/p><p><a href=\"https:\/\/www.davidpfenniglab.com\/wp-content\/uploads\/2020\/09\/Seidl-2019-G3.pdf\">PDF<\/a> Seidl, F., N. A. Levis, R. Schell, D. W. Pfennig, K. S. Pfennig, and I. M. Ehrenreich. 2019. Genome of <em>Spea multiplicata<\/em>, a rapidly developing, phenotypically plastic, and desert-adapted spadefoot toad.<i><b> G3: Genes, Genomes and Genetics <\/b><\/i>3909\u20133919.\u00a0<\/p><p>Seymour, R. S. 1973. Energy metabolism of dormant spadefoot toads (<em>Scaphiopus<\/em>). <i><b>Copeia <\/b><\/i>1973:435\u2013445.<\/p><p>V\u00e1squez, T. & Pfennig, K. S. 2007. Looking on the bright side: females prefer coloration indicative of male size and condition in the sexually dichromatic spadefoot toad, <em>Scaphiopus couchii<\/em>. <strong><em>Behavioral Ecology and Sociobiology<\/em><\/strong> 62: 127-135.<\/p><p>Zeng, C., I. Gomez-Mestre, and J. J. Wiens. 2014. Evolution of rapid development in spadefoot toads is unrelated to arid environments. <i><b>PLoS ONE 9:e96637.<\/b><\/i><\/p>\t\t\t\t\t\t<\/div>\n\t\t\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/div>\n\t\t\t\t\t<\/div>\n\t\t<\/section>\n\t\t\t\t<\/div>\n\t\t","protected":false},"excerpt":{"rendered":"<p>Spadefoots! An overview of their ecology, evolution, development, genetics, and behavior What is a spadefoot? The first thing you need to know about spadefoots toads (hereafter, \u201cspadefoots\u201d) is that they\u2019re not actually toads \u2013\u2013 they\u2019re frogs. True toads are restricted to the genus Bufo (Anaxyrus). \u00a0Spadefoots get their name from the keratinized knob (or \u201cspade\u201d) […]<\/p>\n","protected":false},"author":1,"featured_media":0,"parent":0,"menu_order":0,"comment_status":"closed","ping_status":"closed","template":"","meta":{"footnotes":""},"class_list":["post-25","page","type-page","status-publish","hentry"],"_links":{"self":[{"href":"https:\/\/www.davidpfenniglab.com\/wp-json\/wp\/v2\/pages\/25"}],"collection":[{"href":"https:\/\/www.davidpfenniglab.com\/wp-json\/wp\/v2\/pages"}],"about":[{"href":"https:\/\/www.davidpfenniglab.com\/wp-json\/wp\/v2\/types\/page"}],"author":[{"embeddable":true,"href":"https:\/\/www.davidpfenniglab.com\/wp-json\/wp\/v2\/users\/1"}],"replies":[{"embeddable":true,"href":"https:\/\/www.davidpfenniglab.com\/wp-json\/wp\/v2\/comments?post=25"}],"version-history":[{"count":5,"href":"https:\/\/www.davidpfenniglab.com\/wp-json\/wp\/v2\/pages\/25\/revisions"}],"predecessor-version":[{"id":885,"href":"https:\/\/www.davidpfenniglab.com\/wp-json\/wp\/v2\/pages\/25\/revisions\/885"}],"wp:attachment":[{"href":"https:\/\/www.davidpfenniglab.com\/wp-json\/wp\/v2\/media?parent=25"}],"curies":[{"name":"wp","href":"https:\/\/api.w.org\/{rel}","templated":true}]}}