E. umbellatum population genomics

Errorea gertatu da txantiloia prozesatzerakoan.
The following has evaluated to null or missing:
==> description  [in template "10112#522485#1324421" at line 291, column 15]

----
Tip: If the failing expression is known to legally refer to something that's sometimes null or missing, either specify a default value like myOptionalVar!myDefault, or use <#if myOptionalVar??>when-present<#else>when-missing</#if>. (These only cover the last step of the expression; to cover the whole expression, use parenthesis: (myOptionalVar.foo)!myDefault, (myOptionalVar.foo)??
----

----
FTL stack trace ("~" means nesting-related):
	- Failed at: #if description != ""  [in template "10112#522485#1324421" at line 291, column 9]
----
1<#-- GENERAL--> 
2<#assign cTxtArticle			= languageUtil.get( locale, "article" ) > 
3<#assign cTxtAutoria			= languageUtil.get( locale, "ehu.authors" ) > 
4<#assign cTxtDoi				= languageUtil.get( locale, "ehu.doi" ) > 
5<#assign cTxtDoiTitle			= languageUtil.get( locale, "ehu.digital-object-identifier" ) > 
6<#assign cTxtNewWindow			= languageUtil.get( locale, "opens-new-window" ) > 
7<#assign cTxtPhoto				= languageUtil.get( locale, "ehu.photo" ) > 
8<#assign cTxtPublishing			= languageUtil.get( locale, "publishing" ) > 
9 
10<#assign cUrlDoiOrg				= "https://doi.org/" > 
11<#assign scopeId = themeDisplay.scopeGroupId> 
12<#assign isCampusa = (themeDisplay.getTheme().getContextPath() == "/o/upv-ehu-campusa-theme") > 
13<#assign isGlobal = (themeDisplay.getTheme().getContextPath() == "/o/upv-ehu-global-theme") > 
14<#assign journalLocalService = serviceLocator.findService("com.liferay.journal.service.JournalArticleLocalService")> 
15<#if .vars['reserved-article-id']?has_content> 
16	<#assign articleId = .vars['reserved-article-id'] >  
17<#else> 
18	<#assign articleId = "">  
19</#if> 
20 
21<#if !isGlobal > 
22	<#assign articleClass		= "new" > 
23	<#assign preTitleClass		= "pretitle" > 
24	<#assign titleClass			= "new-title" > 
25	<#assign subTitleClass		= "subtitle" > 
26	<#assign entryClass			= "entry" > 
27	<#assign descriptionClass	= "description" > 
28	<#assign bibliographyClass	= "bibliographic-reference" > 
29	<#assign headerTag			= "div" > 
30	<#assign headerBloqueTag	= "p" > 
31	<#assign htmlHeaderTagClass	= ' class="content-header"' > 
32<#else> 
33	<#assign articleClass		= "new-detail" > 
34	<#assign preTitleClass		= "new-detail__pretitle" > 
35	<#assign titleClass			= "new-detail__title" > 
36	<#assign subTitleClass		= "new-detail__subtitle" > 
37	<#assign entryClass			= "new-detail__body__entry" > 
38	<#assign descriptionClass	= "new-detail__body__description" > 
39	<#assign bibliographyClass	= "new-detail__bibliographic-reference" > 
40	<#assign headerTag			= "header" > 
41	<#assign headerBloqueTag	= "div" > 
42	<#assign htmlHeaderTagClass	= "" > 
43</#if> 
44 
45<#assign entGeneralData = ehugeneraldataFieldSet > 
46 
47<#-- HTML--> 
48<article class="${ articleClass }"> 
49	<#-- PRETITLE, TITLE, SUBTITLE --> 
50	<#assign preTitle = ""> 
51	<#if entGeneralData.ehupretitle?? && entGeneralData.ehupretitle.getData()?? && entGeneralData.ehupretitle.getData()?has_content> 
52		<#assign preTitle = entGeneralData.ehupretitle.getData()>	 
53	</#if> 
54	 
55	<#assign title = ""> 
56	<#if entGeneralData.ehunewtitle?? && entGeneralData.ehunewtitle.getData()?? && entGeneralData.ehunewtitle.getData()?has_content> 
57		<#assign title = entGeneralData.ehunewtitle.getData()> 
58	</#if> 
59	 
60	<#assign subTitle = ""> 
61	<#if entGeneralData.ehunewsubtitle?? && entGeneralData.ehunewsubtitle.getData()?? && entGeneralData.ehunewsubtitle.getData()?has_content> 
62		<#assign subTitle = entGeneralData.ehunewsubtitle.getData()> 
63	</#if> 
64	<${ headerTag }${ htmlHeaderTagClass }> 
65		<#if preTitle != "" > 
66			<${ headerBloqueTag } class="${ preTitleClass }">${ preTitle }</${ headerBloqueTag }> 
67		</#if> 
68		<#if title != "" > 
69			<h1 class="${ titleClass }">${ title }</h1>    
70		</#if> 
71		<#if subTitle != "" > 
72			<${ headerBloqueTag } class="${ subTitleClass }">${ subTitle }</${ headerBloqueTag }> 
73		</#if> 
74	</${ headerTag }> 
75 
76	<#if !isGlobal > 
77		<div class="content-content"> 
78	</#if> 
79	<#if isGlobal > 
80		<div class="new-detail__info-date-share"> 
81	</#if> 
82 
83	<#-- CAMPUSA VOCABULARY --> 
84	<#assign vocabularyName = "campusa" > 
85	<#assign tag_init = "<ul></ul>" > 
86	<#assign categories = tag_init > 
87	<@upvlibs.formatVocabularyCategoriesProperties vocabularyName=vocabularyName /> 
88	<#assign categories = upvlibs.categoriesListStr > 
89	 
90	<#if  categories != tag_init  > 
91		<div class="campusa-category">${ categories }</div> 
92	</#if> 
93 
94	<#-- FIRST PUBLICATION DATE --> 
95	<#if locale == "eu_ES"> 
96		<#assign dateFormat = "yyyy/MM/dd"> 
97	<#else> 
98		<#assign dateFormat = "dd/MM/yyyy"> 
99	</#if> 
100	<#if isGlobal > 
101		<#assign articlePublicationDate = .vars['reserved-article-display-date'].data!"" > 
102		<#if articlePublicationDate != "" > 
103			 
104			<#setting date_format=dateFormat > 
105			<#assign currentLocale = locale> 
106			<#setting locale = localeUtil.getDefault() > 
107			<#if articlePublicationDate?? && articlePublicationDate != "" > 
108				<#assign publicationDate = ( articlePublicationDate?datetime( "EEE, dd MMM yyyy hh:mm:ss" ) )?date > 
109				<#assign publicationDateStr = publicationDate?string > 
110				<div class="new-detail__info-date-share__date"> 
111					<p class="publication_date"> 
112						<strong class="text"> <@liferay.language key="ehu.first-publication-date" />:</strong> 
113					 
114							<span class="date">${ publicationDateStr }</span> 
115					 
116					</p> 
117				</div> 
118			</#if> 
119			<#setting locale = currentLocale> 
120		</#if> 
121	<#else> 
122		<#assign articlePublicationDate = .vars['reserved-article-display-date'].data!"" > 
123		<#if articlePublicationDate != "" >		 
124			<#setting date_format=dateFormat > 
125			<#assign currentLocale = locale> 
126			<#setting locale = localeUtil.getDefault() > 
127			<#if articlePublicationDate?? && articlePublicationDate != "" > 
128				<#assign publicationDate = ( articlePublicationDate?datetime( "EEE, dd MMM yyyy hh:mm:ss" ) )?date > 
129				<#assign publicationDateStr = publicationDate?string > 
130				<p class="publication_date"> 
131					<strong class="text"> <@liferay.language key="ehu.first-publication-date" />:</strong> 
132					<span class="date">${ publicationDateStr }</span> 
133				</p> 
134			</#if> 
135			<#setting locale = currentLocale> 
136		</#if> 
137	</#if> 
138 
139 
140 
141	<#-- REDES SOCIALES -->	 
142		<!-- Si está marcado el check de redes sociales las pintamos --> 
143		<#if entGeneralData.ehusocialbookmarks?? && entGeneralData.ehusocialbookmarks.getData()?has_content && entGeneralData.ehusocialbookmarks.getData()=="true" >		 
144			<#assign classRS = "share-social-bar__item" >			 
145			<#assign urlEnCurso = themeDisplay.getURLPortal() + themeDisplay.getURLCurrent() > 
146			<#-- SHARE SOCIAL BAR --> 
147			<div class="share-social-bar"> 
148				<a href="https://www.facebook.com/share.php?u=${urlEnCurso?replace(' ','%20')}&amp;t=www.ehu.eus" target="_blank" title='<@liferay.language key="ehu.social-bookmark.facebook" />' class="${ classRS } svg-icon"> 
149					<svg xmlns="http://www.w3.org/2000/svg" viewBox="0 0 320 512"><!--!Font Awesome Free 6.7.2 by @fontawesome - https://fontawesome.com License - https://fontawesome.com/license/free Copyright 2025 Fonticons, Inc.--><path d="M80 299.3V512H196V299.3h86.5l18-97.8H196V166.9c0-51.7 20.3-71.5 72.7-71.5c16.3 0 29.4 .4 37 1.2V7.9C291.4 4 256.4 0 236.2 0C129.3 0 80 50.5 80 159.4v42.1H14v97.8H80z"/></svg> 
150					<span class="sr-only"><@liferay.language key="ehu.social-bookmark.facebook" /> - ${ cTxtNewWindow }</span> 
151				</a> 
152				<a href="https://bsky.app/intent/compose?text=${urlEnCurso?replace(' ','%20')}%20-%20www.ehu.eus%20" target="_blank" title='<@liferay.language key="ehu.social-bookmark.bluesky" />' class="${ classRS } svg-icon"> 
153					<svg height="16" viewBox="0 0 512 512" width="16" xmlns="http://www.w3.org/2000/svg"><!--!Font Awesome Free 6.7.2 by @fontawesome - https://fontawesome.com License - https://fontawesome.com/license/free Copyright 2025 Fonticons, Inc.--><path d="M111.8 62.2C170.2 105.9 233 194.7 256 242.4c23-47.6 85.8-136.4 144.2-180.2c42.1-31.6 110.3-56 110.3 21.8c0 15.5-8.9 130.5-14.1 149.2C478.2 298 412 314.6 353.1 304.5c102.9 17.5 129.1 75.5 72.5 133.5c-107.4 110.2-154.3-27.6-166.3-62.9l0 0c-1.7-4.9-2.6-7.8-3.3-7.8s-1.6 3-3.3 7.8l0 0c-12 35.3-59 173.1-166.3 62.9c-56.5-58-30.4-116 72.5-133.5C100 314.6 33.8 298 15.7 233.1C10.4 214.4 1.5 99.4 1.5 83.9c0-77.8 68.2-53.4 110.3-21.8z"></path></svg> 
154					<span class="sr-only"><@liferay.language key="ehu.social-bookmark.bluesky" /> - ${ cTxtNewWindow }</span> 
155				</a> 
156				<a href="https://www.linkedin.com/shareArticle?mini=true&amp;url=${urlEnCurso?replace(' ','%20')}&amp;title=${title?replace(' ','%20')}&amp;summary=" target="_blank" title='<@liferay.language key="ehu.social-bookmark.linkedin" />' class="${ classRS } svg-icon"> 
157					<svg xmlns="http://www.w3.org/2000/svg" viewBox="0 0 448 512"><!--!Font Awesome Free 6.7.2 by @fontawesome - https://fontawesome.com License - https://fontawesome.com/license/free Copyright 2025 Fonticons, Inc.--><path d="M100.3 448H7.4V148.9h92.9zM53.8 108.1C24.1 108.1 0 83.5 0 53.8a53.8 53.8 0 0 1 107.6 0c0 29.7-24.1 54.3-53.8 54.3zM447.9 448h-92.7V302.4c0-34.7-.7-79.2-48.3-79.2-48.3 0-55.7 37.7-55.7 76.7V448h-92.8V148.9h89.1v40.8h1.3c12.4-23.5 42.7-48.3 87.9-48.3 94 0 111.3 61.9 111.3 142.3V448z"/></svg> 
158					<span class="sr-only"><@liferay.language key="ehu.social-bookmark.linkedin" /> - ${ cTxtNewWindow }</span> 
159				</a> 
160				<a href="whatsapp://send?text=${ urlEnCurso }" data-action="share/whatsapp/share" target="_blank" class="${ classRS } svg-icon" title='<@liferay.language key="ehu.social-bookmark.whatsapp" />'> 
161					<svg class="svg-whatsapp" xmlns="http://www.w3.org/2000/svg" viewBox="0 0 448 512"><!--!Font Awesome Free 6.7.2 by @fontawesome - https://fontawesome.com License - https://fontawesome.com/license/free Copyright 2025 Fonticons, Inc.--><path d="M380.9 97.1C339 55.1 283.2 32 223.9 32c-122.4 0-222 99.6-222 222 0 39.1 10.2 77.3 29.6 111L0 480l117.7-30.9c32.4 17.7 68.9 27 106.1 27h.1c122.3 0 224.1-99.6 224.1-222 0-59.3-25.2-115-67.1-157zm-157 341.6c-33.2 0-65.7-8.9-94-25.7l-6.7-4-69.8 18.3L72 359.2l-4.4-7c-18.5-29.4-28.2-63.3-28.2-98.2 0-101.7 82.8-184.5 184.6-184.5 49.3 0 95.6 19.2 130.4 54.1 34.8 34.9 56.2 81.2 56.1 130.5 0 101.8-84.9 184.6-186.6 184.6zm101.2-138.2c-5.5-2.8-32.8-16.2-37.9-18-5.1-1.9-8.8-2.8-12.5 2.8-3.7 5.6-14.3 18-17.6 21.8-3.2 3.7-6.5 4.2-12 1.4-32.6-16.3-54-29.1-75.5-66-5.7-9.8 5.7-9.1 16.3-30.3 1.8-3.7 .9-6.9-.5-9.7-1.4-2.8-12.5-30.1-17.1-41.2-4.5-10.8-9.1-9.3-12.5-9.5-3.2-.2-6.9-.2-10.6-.2-3.7 0-9.7 1.4-14.8 6.9-5.1 5.6-19.4 19-19.4 46.3 0 27.3 19.9 53.7 22.6 57.4 2.8 3.7 39.1 59.7 94.8 83.8 35.2 15.2 49 16.5 66.6 13.9 10.7-1.6 32.8-13.4 37.4-26.4 4.6-13 4.6-24.1 3.2-26.4-1.3-2.5-5-3.9-10.5-6.6z"/></svg>  
162					<span class="sr-only"><@liferay.language key="ehu.social-bookmark.whatsapp" /> - ${ cTxtNewWindow }</span> 
163				</a> 
164				<a href="https://t.me/share/url?url=${ urlEnCurso }&text=${ title }" target="_blank" class="${ classRS } svg-icon" title='<@liferay.language key="ehu.social-bookmark.telegram" />'> 
165					<svg class="svg-telegram" xmlns="http://www.w3.org/2000/svg" viewBox="0 0 496 512"><!--!Font Awesome Free 6.7.2 by @fontawesome - https://fontawesome.com License - https://fontawesome.com/license/free Copyright 2025 Fonticons, Inc.--><path d="M248 8C111 8 0 119 0 256S111 504 248 504 496 393 496 256 385 8 248 8zM363 176.7c-3.7 39.2-19.9 134.4-28.1 178.3-3.5 18.6-10.3 24.8-16.9 25.4-14.4 1.3-25.3-9.5-39.3-18.7-21.8-14.3-34.2-23.2-55.3-37.2-24.5-16.1-8.6-25 5.3-39.5 3.7-3.8 67.1-61.5 68.3-66.7 .2-.7 .3-3.1-1.2-4.4s-3.6-.8-5.1-.5q-3.3 .7-104.6 69.1-14.8 10.2-26.9 9.9c-8.9-.2-25.9-5-38.6-9.1-15.5-5-27.9-7.7-26.8-16.3q.8-6.7 18.5-13.7 108.4-47.2 144.6-62.3c68.9-28.6 83.2-33.6 92.5-33.8 2.1 0 6.6 .5 9.6 2.9a10.5 10.5 0 0 1 3.5 6.7A43.8 43.8 0 0 1 363 176.7z"/></svg>  
166					<span class="sr-only"><@liferay.language key="ehu.social-bookmark.telegram" /> - ${ cTxtNewWindow }</span> 
167				</a> 
168				<a href="mailto:?subject=${title?replace(' ','%20')}&amp;body=%20-%20${urlEnCurso?replace(' ','%20')}" target="_blank" title='<@liferay.language key="ehu.send-email" />' class="${ classRS } svg-icon"> 
169					<svg xmlns="http://www.w3.org/2000/svg" viewBox="0 0 512 512"><!--!Font Awesome Free 6.7.2 by @fontawesome - https://fontawesome.com License - https://fontawesome.com/license/free Copyright 2025 Fonticons, Inc.--><path d="M48 64C21.5 64 0 85.5 0 112c0 15.1 7.1 29.3 19.2 38.4L236.8 313.6c11.4 8.5 27 8.5 38.4 0L492.8 150.4c12.1-9.1 19.2-23.3 19.2-38.4c0-26.5-21.5-48-48-48L48 64zM0 176L0 384c0 35.3 28.7 64 64 64l384 0c35.3 0 64-28.7 64-64l0-208L294.4 339.2c-22.8 17.1-54 17.1-76.8 0L0 176z"/></svg> 
170					<span class="sr-only"><@liferay.language key="ehu.send-email" /> - ${ cTxtNewWindow }</span> 
171				</a> 
172				<a href="javascript:void(0)" onclick="copyCurrentUrl()" title='<@liferay.language key="copy-link" />' class="${ classRS } svg-icon"> 
173					<svg xmlns="http://www.w3.org/2000/svg" viewBox="0 0 640 512"><!--!Font Awesome Free 6.7.2 by @fontawesome - https://fontawesome.com License - https://fontawesome.com/license/free Copyright 2025 Fonticons, Inc.--><path d="M579.8 267.7c56.5-56.5 56.5-148 0-204.5c-50-50-128.8-56.5-186.3-15.4l-1.6 1.1c-14.4 10.3-17.7 30.3-7.4 44.6s30.3 17.7 44.6 7.4l1.6-1.1c32.1-22.9 76-19.3 103.8 8.6c31.5 31.5 31.5 82.5 0 114L422.3 334.8c-31.5 31.5-82.5 31.5-114 0c-27.9-27.9-31.5-71.8-8.6-103.8l1.1-1.6c10.3-14.4 6.9-34.4-7.4-44.6s-34.4-6.9-44.6 7.4l-1.1 1.6C206.5 251.2 213 330 263 380c56.5 56.5 148 56.5 204.5 0L579.8 267.7zM60.2 244.3c-56.5 56.5-56.5 148 0 204.5c50 50 128.8 56.5 186.3 15.4l1.6-1.1c14.4-10.3 17.7-30.3 7.4-44.6s-30.3-17.7-44.6-7.4l-1.6 1.1c-32.1 22.9-76 19.3-103.8-8.6C74 372 74 321 105.5 289.5L217.7 177.2c31.5-31.5 82.5-31.5 114 0c27.9 27.9 31.5 71.8 8.6 103.9l-1.1 1.6c-10.3 14.4-6.9 34.4 7.4 44.6s34.4 6.9 44.6-7.4l1.1-1.6C433.5 260.8 427 182 377 132c-56.5-56.5-148-56.5-204.5 0L60.2 244.3z"/></svg> 
174					<span class="sr-only"><@liferay.language key="copy-link" /> - ${ cTxtNewWindow }</span> 
175				</a> 
176			</div>			 
177		</#if>	 
178 
179	<#if isGlobal > 
180		</div> <#-- class="new-detail__info-date-share" --> 
181		<div class="new-detail__body"> 
182	</#if> 
183 
184	<#-- IMAGE --> 
185	<#assign entImage = entGeneralData.ehunewimageFieldSet > 
186	<#assign imageSrc = "" > 
187	<#if entImage.ehunewimage?? &&  entImage.ehunewimage.getData()?? && entImage.ehunewimage.getData()?has_content> 
188		<#assign imageSrc =  entImage.ehunewimage.getData()> 
189	</#if> 
190	<#if imageSrc != "" > 
191	 
192		<#assign imageAltText = ""> 
193		<#if  entImage.ehunewimagealt?? &&  entImage.ehunewimagealt.getData()?? &&  entImage.ehunewimagealt.getData()?has_content> 
194			<#assign imageAltText = entImage.ehunewimagealt.getData()> 
195		</#if> 
196		 
197		<#assign imageFootText = ""> 
198		<#if entImage.ehunewimagefoot?? &&  entImage.ehunewimagefoot.getData()?? &&  entImage.ehunewimagefoot.getData()?has_content  > 
199			<#assign imageFootText = entImage.ehunewimagefoot.getData() > 
200		</#if> 
201 
202		<#assign imageAuthorText = ""> 
203		<#if entImage.ehunewimageauthor?? &&  entImage.ehunewimageauthor.getData()?? &&  entImage.ehunewimageauthor.getData()?has_content> 
204			<#assign imageAuthorText = entImage.ehunewimageauthor.getData() > 
205		</#if> 
206		 
207		<#assign imageAlign = ""> 
208		<#if entImage.ehuimagedisposition?? && entImage.ehuimagedisposition.getData()?? && entImage.ehuimagedisposition.getData()?has_content> 
209			<#assign imageAlign =  entImage.ehuimagedisposition.getData()> 
210		</#if> 
211		 
212		<#assign imageUrl = "" > 
213		<#if entImage.ehunewimageurl?? && entImage.ehunewimageurl.getData()?? && entImage.ehunewimageurl.getData()?has_content > 
214			<#assign imageUrl = entImage.ehunewimageurl.getData()> 
215		</#if> 
216		 
217	 
218		<#assign imageUrlTitle = "" > 
219		<#assign imageUrlNewTab = true > 
220		<#assign imageClass = "img-main" > 
221 
222		<#-- Pasamos la imagen de la noticia directamente al og:image para que no haya problemas al compartirla 
223			 en redes sociales (Trello 273 - Trello 637) -->	 
224		<@liferay_util["html-top"]> 
225			<meta property="og:image" content="${ portalUtil.getAbsoluteURL( request, imageSrc ) }"> 
226		</@> 
227		<#if !isGlobal && entImage.ehunewimageurl??> 
228			<@upvlibs.imageAuthorSection image=imageSrc altText=imageAltText footText=imageFootText imageAuthor=imageAuthorText 
229				imageDisposition=imageAlign elemImageUrl=entImage.ehunewimageurl imageUrlTitle=imageUrlTitle imageUrlNewTab=imageUrlNewTab imgClass=imageClass /> 
230		<#else> 
231			 
232			<#if imageUrl != ""> 
233				<a href="${ imageUrl }" target="_blank"> 
234			</#if> 
235			<div class="new-detail__body__main-image ${ imageAlign }"> 
236			<img src="${ upvlibs.getImageURL(imageSrc) }" alt="${ imageAltText }" class="${ imageClass }"/>  
237			 
238				<#if imageFootText != "" || imageAuthorText != "" > 
239					<div class="main-image__footer"> 
240						<#if imageFootText == "" > 
241							<#assign txtPhoto = "" > 
242						<#else> 
243							<#assign txtPhoto = imageFootText > 
244						</#if> 
245						<#if imageAuthorText != "" > 
246							<#if imageFootText == "" > 
247								<#assign aux = "" > 
248							<#else> 
249								<#assign aux = " | " > 
250							</#if> 
251							<#assign txtPhoto = txtPhoto + aux + cTxtPhoto + ": " + imageAuthorText > 
252						</#if> 
253						<#if imageAlign == "" > 
254								<#assign classImageFootText = "center" > 
255						<#else> 
256								<#assign classImageFootText = imageAlign > 
257						</#if> 
258						<p class="${ classImageFootText }">${ txtPhoto }</p>					 
259				   </div> 
260				</#if>				 
261		  	 </div> 
262		   <#if imageUrl != ""> 
263				</a> 
264			</#if> 
265		</#if> 
266	<#else> 
267		<#-- Si la imagen no tiene imagen metemos como imagen para redes sociales el logo de la universidad 
268			 proporcionado por la oficina de comunicación (Trello 565 - Trello 637) -->	 
269		<@liferay_util["html-top"]> 
270				<meta property="og:image" content="https://www.ehu.eus/documents/522485/1339603/avatar.jpg"> 
271		</@> 
272	</#if> 
273 
274	<#-- ENTRADILLA --> 
275	<#-- Desde la Oficina de Comunicacion piden para Campusa que la entradilla vaya despues de la imagen 
276		 Se hace el cambio para todos --> 
277	<#assign entradilla = ""> 
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Spatially Explicit Evolution of Diversity (SPEED)

Applicant: Peter B. Pearman (Swiss Federal Research Institute WSL)

Co-applicants: Nicolas Salamin (University of Lausanne), Christian Lexer (University of Freibourgh), Niklaus Zimmermann (Swiss Federal Research Institute), Felix Forest (Royal Botanic Gardens, Kew)

Start date: 1 October 2009

Duration: 44 months

Location of research: South Africa, Switzerland

Support: 964, 637 Swiss Franks

Expanded Abstract:

The species environmental niche consists of the biotic and abiotic conditions necessary for long-term persistence and the niche occupies a central place in the development of ecological theories of competition, limiting similarity, and character divergence.  We study interspecific variation in niche characteristics by adopting an evolutionary perspective.  Application of a toolkit used in the disciplines of systematics and evolutionary biology enables us to address how the environmental niche has changed in a family of plants during the diversification of a clade.  We focus on the Restionaceae, a monophyletic family of grass-like plants of which 340 of the 350 species are endemic to southern Africa.  This is a defensible choice because the relative climatic stability of this region during the Pleistocene suggests that these species distributions are currently at equilibrium with climate.  Under accelerating climate change, the capacity of these species for rapid niche evolution will contribute to determining their fate in a landscape where opportunities for successful dispersal to areas of favourable climate may be limited by intervening expanses of unsuitable habitat and the anthropogenic barriers of agriculture, transportation corridors and urbanization. 

We will develop models of niche evolution that serve both for understanding the evolution of extant species diversity and community structure, and for forecasting how diversity and community structure change, given projected climate change and the potential we identify for rapid niche evolution.  We will model characteristics of the ß-niche (habitat-specific, e.g. geology, fire frequency and vegetation type) and ?-niche (climatic), using field-acquired occurrence data for each species. New GIS data layers of fire frequency and geology will be developed.  The relationship between the fundamental and the estimated realized niche will be experimentally evaluated in multiple species.  We will compare in a greenhouse experiment the drought and flooding tolerances of species to their modelled precipitation requirements.  A transplant experiment in the field will help us determine whether species replacement along an elevation and rainfall gradient is driven primarily by fundamental niche requirements or by interspecific competition and, thus, the realized niche.  We will quantify the evolutionary lability of the environmental niche by modelling selected niche parameters over a completely-sampled species-level phylogeny.  This will indicate which parameters have constrained the evolution of the clade and which parameters are associated with differentiation and speciation.  However, working with species-level phylogenies and niche models has formerly entailed assumptions of constant evolutionary rates and equal inheritance of trait variance by new species.  These assumptions will be addressed in population-level analyses of species complexes to determine speciation modes and establish their impacts on the patterns of inheritance of niche variability.  Further studies will estimate using models of trait evolution the variation among clades in rates of niche change.

Using data on species distributions, chorological analyses and niche modeling we will determine current regional species pools.  These will further be stratified into habitat-specific species pools, using field observations, niche models and models of environmental filtering.  Then, using species traits and phylogenetic relationships, we will seek the determinants of community assembly (i.e., the combinations of species from the regional pool that can be combined into communities).  Using the model parameters developed to address the above questions, we will attempt to estimate the possible contribution of rapid evolution of species environmental niches to change in regional species pools and local communities that may accompany altered climate.  These analyses will provide a powerful tool with which to evaluate and predict the response to climate change by multiple, related species and how these changes could impact regional spatial patterns of diversity in the Restionaceae.

The SPEED project developes the tools necessary to address the effects of climate change and the evolutionary response of species to it.  We intend to develop means to convert knowledge on these changes into geographically-explicit patterns of species diversity in the Restionaceae.  We extend the potential impact of our work by including an integrated program for training three Ph.D. students in the evolutionary and ecological modeling techniques that we develop and use during this research.

Publications:

Lexer, C., R. Wüest, S. Mangili, M. Heuertz, K. N. Stölting, P. B. Pearman, F. Forest, N. Salamin, N. Zimmermann, and E. Bossolini. 2014. Genomics of the speciation continuum in an African plant biodiversity hotspot, I: Drivers of population divergence in Restio capensis (Restionaceae). Molecular Ecology 23:4373-4386.

Litsios, G., R. O. Wüest, A. Kostikova, F. Forest, C. Lexer, H. P. Linder, P. B. Pearman, N. E. Zimmermann and N. Salamin. 2014. Differential effect of fire-survival strategy on diversification among a replicated radiation on two continents. Evolution 68:453-465.

Perret, M., A. Chautems, A. O. De Araujo, and N. Salamin. 2013. Temporal and spatial origin of Gesneriaceae in the New World inferred from plastid DNA sequences. Botanical Journal of the Linnean Society 171:61-79.

Lexer, C., K. Stoelting, S. Mangili, F. Forest, E. Bossolini, P. B. Pearman, N. E. Zimmermann and N. Salamin. 2013. ‘Next generation' biogeography: towards understanding the drivers of species diversification and persistence. Journal of Biogeography 40:1013-1022.

D'Amen, M., N. E. Zimmermann, and P. B. Pearman. 2013. Conservation of phylogeographic lineages under climate change. Global Ecology and Biogeography 22:93-104.

Litsios, G., C. Sims, P. B. Pearman, R. O. Wüest, N. E. Zimmermann, and N. Salamin. 2012. Mutualism with sea anemones triggered the adaptive radiation of clownfish. BMC Evolutionary Biology 12:212.

Litsios, G. and N. Salamin. Effects of phylogenetic signal on ancestral state reconstruction. Systematic Biology 61:533-538.

Litsios, G., L. Pellissier, F. Forest, P. B. Pearman, N. E. Zimmermann, and N. Salamin. 2012. Trophic specialization influences the rate of environmental niche evolution in damselfishes (Pomacentridae). Proc. Royal Soc. B. 279:3662-3669.

Broennimann, O., M. Fitzpatrick, P. B. Pearman B. Petitpierre, L. Pellissier and A. Guisan. 2012.  Measuring ecological niche overlap from occurrence and spatial environmental data. Global Ecology and Biogeography 21: 481-497. 

Pearman, P. B., A. Guisan, and N. E. Zimmermann. 2011. Impacts of climate change on Swiss biodiversity: an indicator species approach. Biological Conservation 144:866-875.

Lexer, C. and K. N. Stölting. 2011. Tracing the recombination and colonization history of hybrid species in space and time. Molecular Ecology 20:3701-3704.

Pearman, P. B., M. D'Amen, C. Graham, W. Thuiller, and N. E. Zimmermann. 2010. Within-taxon niche structure: Niche conservatism, divergence and predicted effects of climate change. Ecography 33:990-1003. (Note: first two authors contributed equally to this paper.)

Salamin, N., R. O. Wüest, S. Lavergne, W. Thuiller and P. B. Pearman. 2010. Assessing rapid evolution in a changing environment. Trends in Ecology and Evolution 25:692-698.
 

Evolutionary Niche Dynamics of Invasive Species

Evolutionary Niche dyNamics of Invasive Species (ENNIS)

Applicant: Peter B. Pearman (Swiss Federal Research Institute WSL)

Co-applicant: Nicolas Salamin (University of Lausanne)

Start date: 1 June 2009

Duration: 41 months

Location of research: California, USA and Switzerland

Support: 240,833 Swiss Franks

Expanded Abstract: 

Ecologists seek to understand the factors that lead some exotic plant species to become ecologically dominant and widespread. This attention is justified because these invasive species can alter community composition, impact ecosystem function, affect species' evolutionary trajectories, and lead to species extinctions. Despite these impacts, finding characteristics that are consistently shared by invasive species is difficult. The climatic niche of plants (the climatic conditions under which population growth is positive) potentially facilitates plant invasions. Some groups of invasive plants have native ranges that span large latitudinal gradients.  Other invasive species have experienced shifts of climatic niche during the invasion process.  Niche shifts, both recent and deep in species evolutionary history, likely influence whether immigrant species become established, naturalized and, subsequently, invasive. The evolutionary dynamics of the climatic niche, per se, have seen very little work and there is substantial disagreement over the occurrence, magnitude and rate of climatic niche shifts. These evolutionary niche dynamics are the focus of this proposal.


The main goal of this project is to understand the influence of evolutionary history, especially the history of shifts of the climatic niche, on the invasiveness of exotic species.  Studying the evolutionary history of niche dynamics is not specific to invasive species. Invasive species are a convenient set of model systems (i.e. genera) in which the literature suggests that it is likely that evolution of the climatic niche influences the presence of a detectable ecological quality: invasiveness. Without the criterion of presence of invasives, we could have selected genera at random or used other criteria that would not likely lead the research to have similarly broad interest. In this research we: (a) use a bioinformatics approach to obtain existing data on species distribution and molecular variation within genera that contain invasive species; (b) modify and use existing software to test the degree to which alternative evolutionary models suffice in describing phylogenetic patterns of niche evolution within these genera; (c) test for evolutionary and historical correlates of niche shift and invasiveness by drawing on information from phylogenetic reconstructions, functional traits, and climatic niche characteristics of species.


The first part of the work focuses on collection of (a) ecological, occurrence, and climate data that describe the distribution-climate relationships of species, (b) data on species functional traits, and (c) existing sequence data for phylogeny reconstructions.  We extract species distribution data from online databases and a wide array of other data sources, including the primary literature. We obtain molecular data directly from GenBank.  Additionally, plant tissue for additional sequencing will be collected at national herbaria and botanical gardens.  We model and quantify niche optima and limits using species distribution models and we construct phylogenetic trees of species.  We model the evolution of the climatic niche using existing open-source software, examining potential effects of both selection and random evolution on niche variability. We produce a supertree of the genera and, using both niche and functional trait data, develop general linear models to examine the relationship of these variables to establishment and invasion by exotic species.  Finally, we expand existing software to examine how heterogeneous selection among subclades affects the likelihood of niche shifts and species invasiveness. 

Beyond the importance of understanding the factors that contribute to species invasiveness, this research will contribute to understanding how niche shifts contribute to evolution within genera and affect species distributions. By using genera that include invasive species in Switzerland and elsewhere, the project will contribute to efforts to identify potentially invasive species before they become introduced.  We will gain better understanding of the role of selection and random evolution in niche shifts of invasive species, closely related species, and their ancestors. By helping us to understand the evolutionary history of niche shifts, this research will improve confidence in the use of species distribution models to predict the potential distributions of invasive species.  Finally, understanding evolutionary processes that enable species to expand into new environments may help to predict more accurately the impacts of climate change on plant biodiversity.

Publications:

Kostikova, A., N. Salamin and P. B. Pearman. 2014. The role of climatic tolerances and seed traits in reduced extinction rates of temperate Polygonaceae. Evolution 68:1856-1870.

Kostikova, A., G. Litsios, S. Burgy, L. Milani, P. B. Pearman, and N. Salamin. 2014. Scale-dependent adaptive evolution and morphological convergence to climate niche in the Californian eriogonoids (Polygonaceae). Journal of Biogeography 41:1326-1337.

Kostikova, A., G. Litsios, N. Salamin, and P. B. Pearman.  2013. Linking life history traits, ecology and niche breadth evolution in the North American eriogonoids (Polygonaceae). American Naturalist 182:760-774.

D'Amen, M., N. E. Zimmermann, and P. B. Pearman. 2013. Conservation of phylogeographic lineages under climate change. Global Ecology and Biogeography 22:93-104.

Broennimann, O., M. Fitzpatrick, P. B. Pearman B. Petitpierre, L. Pellissier and A. Guisan. 2012.  Measuring ecological niche overlap from occurrence and spatial environmental data. Global Ecology and Biogeography 21: 481-497. 

Engler, R. and 19 additional authors.  2011. 21st century climate change threatens mountain flora unequally across Europe. Global Change Biology 17:2330-2341.

Pearman, P. B., A. Guisan, and N. E. Zimmermann. 2011. Impacts of climate change on Swiss biodiversity: an indicator species approach. Biological Conservation 144:866-875.

Pearman, P. B., M. D'Amen, C. Graham, W. Thuiller, and N. E. Zimmermann. 2010. Within-taxon niche structure: Niche conservatism, divergence and predicted effects of climate change. Ecography 33:990-1003.

Salamin, N., R. O. Wüest, S. Lavergne, W. Thuiller and P. B. Pearman. 2010. Assessing rapid evolution in a changing environment. Trends in Ecology and Evolution 25:692-698.