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	<front>
		<journal-meta>
			<journal-id journal-id-type="publisher-id">GRAELL</journal-id>
			<journal-title-group>
				<journal-title>Graellsia</journal-title>
				<abbrev-journal-title abbrev-type="publisher">Graellsia</abbrev-journal-title>
			</journal-title-group>
			<issn publication-format="print">0367-5041</issn>
			<issn publication-format="electronic">1989-953X</issn>
			<issn-l>0367-5041</issn-l>
			<publisher>
				<publisher-name>Consejo Superior de Investigaciones Cient&#x00ED;ficas</publisher-name>
				<publisher-name>SAM</publisher-name>
			</publisher>
		</journal-meta>
		<article-meta>
			<article-id pub-id-type="doi">10.3989/graellsia.2021.v77.324</article-id>
			<article-id pub-id-type="publisher-id">graellsia.2021.v77.324</article-id>
			<article-categories>
				<subj-group subj-group-type="heading">
					<subject>Art&#x00ED;culo</subject>
				</subj-group>
			</article-categories>
			<title-group>
				<article-title>A new species of <italic>Zwicknia</italic> Mur&#x00E1;nyi, 2014 from Spain (Plecoptera: Capniidae)</article-title>
				<trans-title-group xml:lang="es">
					<trans-title>Una nueva especie de <italic>Zwicknia</italic> Mur&#x00E1;nyi, 2014 de Espa&#x00F1;a (Plecoptera: Capniidae)</trans-title>
				</trans-title-group>
				<alt-title alt-title-type="running-head">New species of <italic>Zwicknia</italic> from Spain</alt-title>
			</title-group>
			<contrib-group>
				<contrib contrib-type="author" corresp="yes">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0001-9424-9576</contrib-id>
					<name name-style="western">
						<surname>Reding</surname>
						<given-names>Jean-Paul G.</given-names>
					</name>
					<xref ref-type="aff" rid="aff-1-e150">1</xref>
					<aff id="aff-1-e150">
						<label>1</label>
						<addr-line>Petit-Berne 2, CH-2035 Corcelles</addr-line>, <country>Switzerland</country>
					</aff>
					<email xlink:href="jean-paul.reding@bluewin.ch">jean-paul.reding@bluewin.ch</email>
				</contrib>
				<contrib contrib-type="author">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-4553-1310</contrib-id>
					<name name-style="western">
						<surname>Pardo</surname>
						<given-names>Isabel</given-names>
					</name>
					<xref ref-type="aff" rid="aff-2-e150">2</xref>
					<aff id="aff-2-e150">
						<label>2</label>
						<institution>Department of Ecology and Animal Biology, University of Vigo</institution>, <addr-line>36310 Vigo</addr-line>, <country>Spain</country>
					</aff>
					<email xlink:href="ipardo@uvigo.es">ipardo@uvigo.es</email>
				</contrib>
				<contrib contrib-type="author">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0003-4702-5136</contrib-id>
					<name name-style="western">
						<surname>Vin&#x00E7;on</surname>
						<given-names>Gilles</given-names>
					</name>
					<xref ref-type="aff" rid="aff-3-e150">3</xref>
					<aff id="aff-3-e150">
						<label>3</label>
						<addr-line>Bd Joseph Vallier 55, F-38100 Grenoble</addr-line>, <country>France</country>
					</aff>
					<email xlink:href="gvincon@gmail.com">gvincon@gmail.com</email>
				</contrib>
			</contrib-group>
			<pub-date pub-type="epub">
				<day>XX</day>
				<month>XX</month>
				<year>2021</year>
			</pub-date>
			<pub-date pub-type="collection">
				<month>12</month>
				<year>2021</year>
			</pub-date>
			<volume>77</volume>
			<issue>2</issue>
			<elocation-id>e150</elocation-id>
			<history>
				<date date-type="received">
					<day>04</day>
					<month>08</month>
					<year>2021</year>
				</date>
				<date date-type="accepted">
					<day>10</day>
					<month>09</month>
					<year>2021</year>
				</date>
				<date date-type="available-online">
					<day>02</day>
					<month>12</month>
					<year>2021</year>
				</date>
			</history>
			<permissions>
				<copyright-statement>&#x00A9; 2021 SAM &#x0026; CSIC</copyright-statement>
				<copyright-year>2021</copyright-year>
				<license license-type="open-access" xlink:href="https://creativecommons.org/licenses/by/4.0/">
					<license-p>This is an open-access article distributed under the terms of the Creative Commons Attribution 4.0 International (CC BY 4.0) License.</license-p>
				</license>
			</permissions>
			<self-uri xlink:href="GRAELL-77-2-e150.pdf"/>
			<abstract>
				<p>A new species, <italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold>, from Spain is described and illustrated from male and female adults and last instar larvae. Morphological affinities of this new species are also discussed and a key to adult <italic>Zwicknia</italic>
					<xref ref-type="bibr" rid="ref-14-e150">Mur&#x00E1;nyi, 2014</xref> from Western and Southern Europe is proposed. From the morphological point of view <italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold> clearly differs from all other known species or subspecies of the <italic>Zwicknia</italic> genus.</p>
					<p><ext-link ext-link-type="uri" xlink:href="http://zoobank.org/urn:lsid:zoobank.org:pub:1386DF7A-DEB0-46A7-B48E-FE19C5329F6D">urn:lsid:zoobank.org:pub:1386DF7A-DEB0-46A7-B48E-FE19C5329F6D</ext-link></p>
			</abstract>
			<trans-abstract xml:lang="es">
				<p>Se describe e ilustra una nueva especie, <italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold>, de Espa&#x00F1;a a partir de machos y hembras adultos y de larvas del &#x00FA;ltimo estadio. Tambi&#x00E9;n se discuten las afinidades morfol&#x00F3;gicas de esta nueva especie y se propone una clave para los adultos de <italic>Zwicknia</italic> de Europa occidental y meridional. Desde el punto de vista morfol&#x00F3;gico, <italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold> difiere claramente de todas las dem&#x00E1;s especies o subespecies conocidas del g&#x00E9;nero <italic>Zwicknia</italic>.</p>
			</trans-abstract>
			<kwd-group>
				<kwd>
					<italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold>
				</kwd>
				<kwd>Spain</kwd>
				<kwd>alpha taxonomy</kwd>
				<kwd>identification key</kwd>
			</kwd-group>
			<kwd-group xml:lang="es">
				<kwd>
					<italic>Zwicknia avilensis</italic>
					<bold>n. sp.</bold>
				</kwd>
				<kwd>Espa&#x00F1;a</kwd>
				<kwd>taxonom&#x00ED;a alfa</kwd>
				<kwd>clave de identificaci&#x00F3;n</kwd>
			</kwd-group>
			<counts>
				<page-count count="14"/>
			</counts>
		</article-meta>
	</front>
	<body>
		<sec sec-type="intro" id="sec-1-e150">
			<title>Introduction</title>
			<p>The establishment of the genus <italic>Zwicknia</italic>
				<xref ref-type="bibr" rid="ref-14-e150">Mur&#x00E1;nyi, 2014</xref> and the partial revision of the genus <italic>Capnia</italic> Pictet, 1841 by <xref ref-type="bibr" rid="ref-14-e150">Mur&#x00E1;nyi <italic>et al.</italic> (2014)</xref> have induced several important changes in the taxonomy of the Capniidae, thus rendering necessary to revise the material collected in Spain under these labels. In this region, the family Capniidae is represented by the four genera <italic>Capnia</italic>, <italic>Capnioneura</italic> Ris, 1905, <italic>Capnopsis</italic> Morton, 1896 and <italic>Zwicknia</italic>. Whereas the status of <italic>Capnioneura</italic> (six species, <italic>Capnioneura petitpierreae</italic> Aubert, 1961, <italic>C. brachyptera</italic> Despax, 1932b, <italic>C. libera</italic> (Nav&#x00E1;s, 1909), <italic>C. mitis</italic> Despax, 1932a, <italic>C. gelesae</italic> Berth&#x00E9;lemy &#x0026; Baena, 1984, <italic>C. narcea</italic> Vin&#x00E7;on &#x0026; S&#x00E1;nchez-Ortega, 2002) and <italic>Capnopsis</italic> (one species, <italic>C. schilleri</italic> (Rostock, 1892)) remain unchanged in Spain (<xref ref-type="bibr" rid="ref-22-e150">Tierno de Figueroa <italic>et al.</italic>, 2003</xref>, <xref ref-type="bibr" rid="ref-23-e150">2014</xref>, <xref ref-type="bibr" rid="ref-24-e150">2018</xref>), the former genus <italic>Capnia</italic> is now subdivided into three different taxa: a genus <italic>Capnia sensu stricto, sensu</italic>
				<xref ref-type="bibr" rid="ref-14-e150">Mur&#x00E1;nyi, 2014</xref>, which has as its sole representative the widely distributed transpalearctic <italic>Capnia nigra</italic> (Pictet, 1833), then a genus <italic>Capnia sensu lato, sensu</italic>
				<xref ref-type="bibr" rid="ref-14-e150">Mur&#x00E1;nyi, 2014</xref>, which includes only the species <italic>Capnia vidua</italic> Klap&#x00E1;lek, 1904 (<xref ref-type="bibr" rid="ref-13-e150">Luz&#x00F3;n-Ortega <italic>et al.</italic>, 2013</xref>; <xref ref-type="bibr" rid="ref-23-e150">Tierno de Figueroa <italic>et al.</italic>, 2014</xref>), and finally, the genus <italic>Zwicknia</italic>, which now replaces the former species complex <italic>Capnia bifrons</italic> (Newman, 1838). Upon examination, it turned out that specimens of <italic>Zwicknia</italic> from the Province of &#x00C1;vila, Spain, that had previously (<xref ref-type="bibr" rid="ref-9-e150">Gonz&#x00E1;lez del T&#x00E1;nago, 1984</xref>) been identified as the now obsolete taxon <italic>Capnia bifrons</italic>, could not be assigned to one of the already known twelve species or subspecies of <italic>Zwicknia</italic> (<xref ref-type="bibr" rid="ref-8-e150">DeWalt <italic>et al.</italic>, 2021</xref>) and gave rise therefore to the suspicion that they corresponded to an unknown taxon. Fresh material collected from the same sampling station in winter 2020 and spring 2021 by the second author have confirmed these initial findings, enabling the description of this new species in the present contribution.</p>
		</sec>
		<sec sec-type="materialsmethods" id="sec-2-e150">
			<title>Material and methods</title>
			<p>C<sc>ollecting</sc>. Adults of <italic>Zwicknia</italic> were collected by using a &#x201C;Japanese umbrella&#x201D; (beating sheet). Nymphs of <italic>Zwicknia</italic>, together with other aquatic macroinvertebrates, were collected by using a hand net and by removing the substratum (<xref ref-type="bibr" rid="ref-9-e150">Gonz&#x00E1;lez del T&#x00E1;nago, 1984</xref>). The older material studied in this contribution had been collected by Dr. Marta Gonz&#x00E1;lez del T&#x00E1;nago in 1981. These specimens were initially stored in the private collection of Dr. Marta Gonz&#x00E1;lez del T&#x00E1;nago, and later, in the 1990s, transferred to the A. S&#x00E1;nchez-Ortega collection in the Department of Zoology (formerly Department of Animal Biology and Ecology) of the University of Granada (Spain). Additional adult specimens have been collected by the second author in winter 2020 and spring 2021. The material is preserved in ethanol.</p>
			<p>T<sc>erminology and abbreviations</sc>. Terminology for adults follows that of <xref ref-type="bibr" rid="ref-14-e150">Mur&#x00E1;nyi <italic>et al.</italic> (2014)</xref>. The wing venation terminology used within this paper follows <xref ref-type="bibr" rid="ref-5-e150">B&#x00E9;thoux (2005)</xref> and <xref ref-type="bibr" rid="ref-6-e150">B&#x00E9;thoux <italic>et al.</italic> (2015)</xref>. The following abbreviations are used: m = adult males; f = adult females; Lm = male larvae; Lf = female larvae; MZL = collections of the Zoological Museum of Lausanne, Switzerland; GVC = collection of G. Vin&#x00E7;on; RC = collection of J.-P.G. Reding; SOC = collection of S&#x00E1;nchez-Ortega, Department of Zoology, University of Granada, Spain.</p>
			<p>M<sc>orphology and depositories</sc>. For the identification of adults and larvae, <xref ref-type="bibr" rid="ref-14-e150">Mur&#x00E1;nyi <italic>et al.</italic> (2014)</xref>, <xref ref-type="bibr" rid="ref-7-e150">Boumans &#x0026; Mur&#x00E1;nyi (2014)</xref>, <xref ref-type="bibr" rid="ref-19-e150">Reding <italic>et al.</italic> (2016)</xref>, <xref ref-type="bibr" rid="ref-17-e150">Reding (2020</xref>, <xref ref-type="bibr" rid="ref-18-e150">2021</xref>) and <xref ref-type="bibr" rid="ref-26-e150">Vin&#x00E7;on &#x0026; Reding (2018)</xref> references were used. Adults of the following six species from Western and Southern Europe were examined for comparison: <italic>Zwicknia acuta</italic>
				<xref ref-type="bibr" rid="ref-14-e150">Mur&#x00E1;nyi &#x0026; Orci, 2014</xref>, <italic>Z. bifrons</italic>, <italic>Z. gattolliati</italic>
				<xref ref-type="bibr" rid="ref-26-e150">Vin&#x00E7;on &#x0026; Reding, 2018</xref>, <italic>Z. ledoarei</italic>
				<xref ref-type="bibr" rid="ref-19-e150">Reding, Launay, Ruffoni, Vin&#x00E7;on &#x0026; Boumans, 2016</xref>, <italic>Z. rupprechti</italic>
				<xref ref-type="bibr" rid="ref-14-e150">Mur&#x00E1;nyi, Orci &#x0026; Gamboa, 2014</xref> and <italic>Z. westermanni</italic>
				<xref ref-type="bibr" rid="ref-7-e150">Boumans &#x0026; Mur&#x00E1;nyi, 2014</xref>. Illustrations of the specimens were produced by the first author with the help of a Nikon E5400 camera attached to a Leica S8-APO stereomicroscope. The photograph of the sampling station was taken by second author. Holotype and paratypes are deposited in the Zoological Museum of Lausanne, Switzerland.</p>
		</sec>
		<sec sec-type="results" id="sec-3-e150">
			<title>Results</title>
			<p>
				<bold>
					<italic>Zwicknia avilensis</italic>
				</bold> sp. n.</p>
				<p><ext-link ext-link-type="uri" xlink:href="http://zoobank.org/urn:lsid:zoobank.org:act:25186B8C-A757-4ED7-A67A-062A70575A10">urn:lsid:zoobank.org:act:25186B8C-A757-4ED7-A67A-062A70575A10</ext-link></p>
			<p>
				<xref ref-type="fig" rid="fig-1-e150">Figs. 1</xref>&#x2013;<xref ref-type="fig" rid="fig-3-e150">13</xref>
			</p>
			<fig id="fig-1-e150">
				<label>Figs. 1&#x2013;6</label>
				<caption>
					<title>
						<italic>Zwicknia avilensis</italic>
						<bold>sp. n.</bold>, male. <bold>1</bold>, epiproct and process on tergite 9, lateral view; <bold>2</bold>, epiproct and process on tergite 9, lateral view; <bold>3</bold>, epiproct and process on tergite 9, lateral view; <bold>4</bold>, epiproct and process on tergite 9, lateral view (&#x00AB;hunchback&#x00BB; marked by arrow); <bold>5</bold>, process on tergite 9, plain dorsal view; <bold>6</bold>, process on tergite 9, caudal view.</title>
					<p xml:lang="es">Figs. 1&#x2013;6.&#x2014; <italic>Zwicknia avilensis</italic>
						<bold>sp. n.</bold>, macho. <bold>1</bold>, epiprocto y proceso del 9º terguito, vista lateral; <bold>2</bold>, epiprocto y proceso del 9º terguito, vista lateral; <bold>3</bold>, epiprocto y proceso del 9º terguito, vista lateral; <bold>4</bold>, epiprocto y proceso del 9º terguito, vista lateral (&#x00AB;joroba&#x00BB; marcada con flecha); <bold>5</bold>, proceso del 9º terguito, vista dorsal; <bold>6</bold>, proceso del 9º terguito, vista caudal.</p>
				</caption>
				<graphic id="gra-1-e150" xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="GRAELL-77-2-e150-f1-G.png"/>
			</fig>
			<p>
				<italic>Capnia bifrons</italic> (Newman, 1838) &#x2013; <xref ref-type="bibr" rid="ref-9-e150">Gonz&#x00E1;lez del T&#x00E1;nago, 1984: 53</xref>.</p>
			<p>T<sc>ype material</sc>. <bold>Holotype male</bold>: SPAIN, &#x00C1;vila Province (05143), Sierra de la Serrota, Arroyo de Canto Moreno, Mu&#x00F1;otello, Duero basin; 40&#x00B0;3337.13 N, 5&#x00B0; 0129.02 E, 1129 m a.s.l., 26.XII.2020, leg. Isabel Pardo, deposited in the MZL (catalogue number: GBIFCH00830015).</p>
			<p>
				<bold>Paratypes:</bold> same locality (<xref ref-type="fig" rid="fig-11-e150">Fig. 56</xref>), same date, 1m, 1f, leg. Isabel Pardo, deposited in the MZL (catalogue numbers: GBIFCH00830016 (male); GBIFCH00830017 (female).</p>
			<p>
				<bold>Additional specimens</bold> are held in the collections of SOC, GVC and RC: <bold>SPAIN:</bold> &#x00C1;vila Province (05143), Sierra de la Serrota, Arroyo de Canto Moreno, Mu&#x00F1;otello, Duero basin; 40&#x00B0;3337.13 N, 5&#x00B0; 0129.02 E, 1129 m; 15.III.1981, 3m, 3f (leg. Marta Gonz&#x00E1;lez del T&#x00E1;nago; GVC); 15.III.1981, 3m, 2f; 6Lm, 6Lf (leg. Marta Gonz&#x00E1;lez del T&#x00E1;nago; SOC); 02.IV.2021, 2f (leg. Isabel Pardo; 1f, RC; 1f, MZL used for molecular studies (catalogue number: GBIFCH00829872).</p>
			<sec id="sec-3.1-e150">
				<title>M<sc>orphological diagnosis</sc></title>
				<p>Male strongly micropterous; female macropterous. Process of male tergite 9 projecting headward (<xref ref-type="fig" rid="fig-1-e150">Figs. 1&#x2013;4</xref>, <xref ref-type="fig" rid="fig-2-e150">7&#x2013;8</xref>). Main epiproct sclerite slender, smoothly truncated and with a slightly blunted tip in lateral view (<xref ref-type="fig" rid="fig-1-e150">Figs. 2&#x2013;3</xref>). Main epiproct sclerite with a small &#x201C;hunchback&#x201D; dorsally in its middle section (<xref ref-type="fig" rid="fig-1-e150">Figs. 3&#x2013;4</xref>).</p>
			</sec>
			<sec id="sec-3.2-e150">
				<title>D<sc>escription</sc></title>
				<p>Head, thorax, appendages and basal segments of the abdomen as typical for the genus. Male strongly micropterous (<xref ref-type="fig" rid="fig-2-e150">Fig. 9</xref>). Female macropterous. Body length: holotype 5.0 mm, male paratypes 4.9&#x2013;7.5 mm, female paratypes 9.0&#x2013;10.1 mm; forewing length of males: holotype 1.1 mm, male paratypes 0.5&#x2013;1.8 mm, female paratypes 10.0&#x2013;11.0 mm.</p>
				<fig id="fig-2-e150">
					<label>Figs. 7&#x2013;12</label>
					<caption>
						<title>
							<bold>7&#x2013;10</bold>. <italic>Zwicknia avilensis</italic>
							<bold>sp. n.</bold>, male. <bold>7</bold>, process on tergite 9, plain dorsal view; <bold>8</bold>, process on tergite 9, ¾ dorsal view; <bold>9</bold>, wings; <bold>10</bold>, ventral vesicle. <bold>11&#x2013;12.</bold>
							<italic>Zwicknia avilensis</italic>
							<bold>sp. n.</bold>, female. <bold>11</bold>, femur of hindleg, lateral view; <bold>12</bold>, subgenital plate, ventral view.</title>
						<p xml:lang="es">Figs. 7&#x2013;12.&#x2014; <bold>7&#x2013;10</bold>. <italic>Zwicknia avilensis</italic>
							<bold>sp. n.</bold>, macho. <bold>7</bold>, proceso del 9º terguito, vista dorsal, <bold>8</bold>, proceso del 9º terguito, ¾ vista dorsal; <bold>9</bold>, alas; <bold>10</bold>, ves&#x00ED;cula ventral. <bold>11&#x2013;12.</bold>
							<italic>Zwicknia avilensis</italic>
							<bold>sp. n.</bold>, hembra. <bold>11</bold>, f&#x00E9;mur posterior, vista lateral; <bold>12</bold>, placa subgenital, vista ventral.</p>
					</caption>
					<graphic id="gra-2-e150" xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="GRAELL-77-2-e150-f2-G.png"/>
				</fig>
				<p>
					<italic>Males</italic> (<xref ref-type="fig" rid="fig-1-e150">Figs. 1</xref>&#x2013;<xref ref-type="fig" rid="fig-2-e150">10</xref>). Process of tergite 9 highly elevated and projecting headward, in lateral (<xref ref-type="fig" rid="fig-1-e150">Figs. 1&#x2013;4</xref>) and dorsal (<xref ref-type="fig" rid="fig-2-e150">Figs, 7&#x2013;8</xref>) views. In caudal view, process of tergite 9 with a wide base and a dome-shaped apex (<xref ref-type="fig" rid="fig-1-e150">Figs. 6</xref>&#x2013;<xref ref-type="fig" rid="fig-2-e150">8</xref>). In lateral view, the main epiproct sclerite is long and slender, with a slightly blunted tip, neither upcurved nor downcurved (<xref ref-type="fig" rid="fig-1-e150">Figs. 2&#x2013;3</xref>). Dorsal part of apex of epiproct smoothly truncated, in lateral view (<xref ref-type="fig" rid="fig-1-e150">Figs. 1&#x2013;4</xref>); ventral part straight. Main epiproct sclerite with a small &#x201C;hunchback&#x201D; in its middle section (<xref ref-type="fig" rid="fig-1-e150">Figs. 3&#x2013;4</xref>, arrow). Membranous part of the apex of the main epiproct sclerite very long, in lateral view (<xref ref-type="fig" rid="fig-1-e150">Figs. 3&#x2013;4</xref>). In dorso-caudal view, the main epiproct sclerite is long and slender, with nearly parallel edges and not swollen medially (<xref ref-type="fig" rid="fig-1-e150">Figs. 5</xref>, <xref ref-type="fig" rid="fig-2-e150">7&#x2013;8</xref>). Apex of process on tergite 9 at least three times as wide as the tip of the main epiproct sclerite, in dorsal view (<xref ref-type="fig" rid="fig-2-e150">Figs. 7&#x2013;8</xref>). Ventral vesicle very large and rounded, its size corresponding to one third of the width of the subgenital plate (<xref ref-type="fig" rid="fig-2-e150">Fig. 10</xref>). Subgenital plate heart-shaped, as typical for the genus (<xref ref-type="fig" rid="fig-2-e150">Fig. 10</xref>).</p>
				<p>
					<italic>Females</italic> (<xref ref-type="fig" rid="fig-2-e150">Figs. 11</xref>&#x2013;<xref ref-type="fig" rid="fig-3-e150">13</xref>). Wing venation as typical for the genus, with the first cell between CuA and CuP in anterior wings quadrangular in shape (<xref ref-type="fig" rid="fig-3-e150">Fig. 13</xref>; cf. <xref ref-type="fig" rid="fig-10-e150">Fig. 55</xref>; <xref ref-type="bibr" rid="ref-4-e150">Aubert 1959: 26, fig. 22</xref>; <xref ref-type="bibr" rid="ref-17-e150">Reding 2020: Vol. 4, fig. 17A20</xref>) and not triangular, as in <italic>Capnia nigra</italic> (<xref ref-type="fig" rid="fig-3-e150">Fig. 14</xref>) or <italic>Capnia vidua</italic>. Dorsal edge of femora and tibiae of all three legs with short bristles (<xref ref-type="fig" rid="fig-2-e150">Fig. 11</xref>). Posterior edge of female subgenital plate crescent-shaped (<xref ref-type="fig" rid="fig-2-e150">Fig. 12</xref>).</p>
				<fig id="fig-3-e150">
					<label>Figs. 13&#x2013;14</label>
					<caption>
						<title>
							<bold>13</bold>. <italic>Zwicknia avilensis</italic>
							<bold>sp. n.</bold>, female, wing venation of anterior wing. <bold>14</bold>. <italic>Capnia nigra</italic>, wing venation of anterior wing of adult female (annotations by Dr. Olivier B&#x00E9;thoux).</title>
						<p xml:lang="es">Figs. 13&#x2013;14.&#x2014; <bold>13</bold>. <italic>Zwicknia avilensis</italic>
							<bold>sp. n.</bold>, hembra, venaci&#x00F3;n del ala anterior. <bold>14</bold>. <italic>Capnia nigra</italic>, venaci&#x00F3;n del ala anterior de la hembra adulta (anotaciones del Dr. Olivier B&#x00E9;thoux).</p>
					</caption>
					<graphic id="gra-3-e150" xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="GRAELL-77-2-e150-f3-G.png"/>
				</fig>
				<fig id="fig-4-e150">
					<label>Figs. 15&#x2013;20</label>
					<caption>
						<title>
							<italic>Zwicknia avilensis</italic>
							<bold>sp. n.</bold>, larva. <bold>15</bold>, bristles on head, lateral view; <bold>16</bold>, bristles on pronotum, lateral view; <bold>17</bold>, bristles on head and pronotum, lateral view; <bold>18</bold>, tergites, dorsal view; <bold>19</bold>, tergites, dorso-lateral view; <bold>20</bold>, dorsal edge of femur of hindleg, lateral view.</title>
						<p xml:lang="es">Figs. 15&#x2013;20.&#x2014; <italic>Zwicknia avilensis</italic>
							<bold>sp. n.</bold>, larva. <bold>15</bold>, cerdas de la cabeza, vista lateral; <bold>16</bold>, cerdas del pronoto, vista lateral; <bold>17</bold>, cerdas de la cabeza y pronoto, vista lateral; <bold>18</bold>, terguitos, vista dorsal; <bold>19</bold>, terguitos, vista dorso-lateral; <bold>20</bold>, borde dorsal del f&#x00E9;mur posterior, vista lateral.</p>
					</caption>
					<graphic id="gra-4-e150" xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="GRAELL-77-2-e150-f4-G.png"/>
				</fig>
				<fig id="fig-5-e150">
					<label>Figs. 21&#x2013;26</label>
					<caption>
						<title>
							<italic>Zwicknia avilensis</italic>
							<bold>sp. n.</bold>, larva. <bold>21</bold>, dorsal edge of tibia of hindleg of male larva, lateral view; <bold>22</bold>, dorsal edge of tibia of hindleg of female larva, lateral view; <bold>23</bold>, micropterism prefigured on male larva, dorsal view; <bold>24</bold>, epiproct and process of tergite 9 of mature male larva, dorsal view; <bold>25</bold>, epiproct and process of tergite 9 of near mature male larva, lateral view; <bold>26</bold>, epiproct and process of tergite 9 of mature male larva, lateral view.</title>
						<p xml:lang="es">Figs. 21&#x2013;26.&#x2014; <italic>Zwicknia avilensis</italic>
							<bold>sp. n.</bold>, larva. <bold>21</bold>, borde dorsal de la tibia posterior de la larva macho, vista lateral; <bold>22</bold>, borde dorsal de la tibia posterior de la larva hembra, vista lateral; <bold>23</bold>, micropterismo prefigurado en la larva macho, vista dorsal; <bold>24</bold>, epiprocto y proceso del 9º terguito de la larva macho madura, vista dorsal; <bold>25</bold>, epiprocto y proceso del 9º terguito de la larva macho casi madura, vista lateral; <bold>26</bold>, epiprocto y proceso del 9º terguito de la larva macho madura, vista lateral.</p>
					</caption>
					<graphic id="gra-5-e150" xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="GRAELL-77-2-e150-f5-G.png"/>
				</fig>
				<fig id="fig-6-e150">
					<label>Figs. 27&#x2013;32</label>
					<caption>
						<title>Epiproct and process on tergite 9 of adult males of <italic>Zwicknia</italic>. <bold>27</bold>, <italic>Zwicknia bifrons</italic> (Jura Mountains); <bold>28</bold>, <italic>Zwicknia acuta</italic> (Slovakia); <bold>29</bold>, <italic>Zwicknia westermanni</italic> (Occitanie); <bold>30</bold>, <italic>Zwicknia gattolliati</italic> (Apennines); <bold>31</bold>, <italic>Zwicknia ledoarei</italic> (Jura Mountains); <bold>32</bold>, <italic>Zwicknia rupprechti</italic> (Vosges).</title>
						<p xml:lang="es">Figs. 27&#x2013;32.&#x2014; Epiprocto y proceso del 9º terguito de machos adultos de <italic>Zwicknia.</italic>
							<bold>27</bold>, <italic>Zwicknia bifrons</italic> (Monta&#x00F1;as del Jura); <bold>28</bold>, <italic>Zwicknia acuta</italic> (Eslovaquia); <bold>29</bold>, <italic>Zwicknia westermanni</italic> (Occitania); <bold>30</bold>, <italic>Zwicknia gattolliati</italic> (Apeninos); <bold>31</bold>, <italic>Zwicknia ledoarei</italic> (Monta&#x00F1;as del Jura); <bold>32</bold>, <italic>Zwicknia rupprechti</italic> (Vosgos).</p>
					</caption>
					<graphic id="gra-6-e150" xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="GRAELL-77-2-e150-f6-G.png"/>
				</fig>
				<fig id="fig-7-e150">
					<label>Figs. 33&#x2013;38</label>
					<caption>
						<title>Ventral vesicle of adult males of <italic>Zwicknia.</italic>
							<bold>33</bold>, <italic>Zwicknia bifrons</italic> (Jura Mountains); <bold>34</bold>, <italic>Zwicknia acuta</italic> (Slovakia); <bold>35</bold>, <italic>Zwicknia westermanni</italic> (Occitanie); <bold>36</bold>, <italic>Zwicknia gattolliati</italic> (Apennines); <bold>37</bold>, <italic>Zwicknia ledoarei</italic> (Jura Mountains); <bold>38</bold>, <italic>Zwicknia rupprechti</italic> (Vosges).</title>
						<p xml:lang="es">Figs. 33&#x2013;38.&#x2014; Ves&#x00ED;cula ventral de machos adultos de <italic>Zwicknia</italic>. <bold>33</bold>, <italic>Zwicknia bifrons</italic> (Monta&#x00F1;as del Jura); <bold>34</bold>, <italic>Zwicknia acuta</italic> (Eslovaquia); <bold>35</bold>, <italic>Zwicknia westermanni</italic> (Occitania); <bold>36</bold>, <italic>Zwicknia gattolliati</italic> (Apeninos); <bold>37</bold>, <italic>Zwicknia ledoarei</italic> (Monta&#x00F1;as del Jura); <bold>38</bold>, <italic>Zwicknia rupprechti</italic> (Vosgos).</p>
					</caption>
					<graphic id="gra-7-e150" xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="GRAELL-77-2-e150-f7-G.png"/>
				</fig>
				<fig id="fig-8-e150">
					<label>Figs. 39&#x2013;44</label>
					<caption>
						<title>Adult females of <italic>Zwicknia</italic>. <bold>39</bold>, femur of hindleg of <italic>Zwicknia bifrons</italic> (Jura Mountains); <bold>40</bold>, femur of hindleg of <italic>Zwicknia gattolliati</italic> (Apennines); <bold>41</bold>, subgenital plate of <italic>Zwicknia bifrons</italic> (Jura Mountains); <bold>42</bold>, subgenital plate of <italic>Zwicknia ledoarei</italic> (Jura Mountains); <bold>43</bold>, sternites and subgenital plate of <italic>Zwicknia gattolliati</italic> (Apennines); <bold>44</bold>, subgenital plate of <italic>Zwicknia westermanni</italic> (Occitanie).</title>
						<p xml:lang="es">Figs. 39&#x2013;44.&#x2014; Hembras adultas de <italic>Zwicknia</italic>. <bold>39</bold>, femur posterior de <italic>Zwicknia bifrons</italic> (Monta&#x00F1;as del Jura); <bold>40</bold>, femur posterior de <italic>Zwicknia gattolliati</italic> (Apeninos); <bold>41</bold>, placa subgenital de <italic>Zwicknia bifrons</italic> (Monta&#x00F1;as del Jura); <bold>42</bold>, placa subgenital de <italic>Zwicknia ledoarei</italic> (Monta&#x00F1;as del Jura); <bold>43</bold>, esternitos y placa subgenital de <italic>Zwicknia gattolliati</italic> (Apeninos); <bold>44</bold>, placa subgenital de <italic>Zwicknia westermanni</italic> (Occitania).</p>
					</caption>
					<graphic id="gra-8-e150" xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="GRAELL-77-2-e150-f8-G.png"/>
				</fig>
				<fig id="fig-9-e150">
					<label>Figs. 45&#x2013;52</label>
					<caption>
						<title>
							<bold>45&#x2013;51</bold>. Lateral view of epiproct of mature male larvae of <italic>Capnia</italic> and <italic>Zwicknia.</italic>
							<bold>45</bold>, <italic>Capnia vidua</italic> (Swiss Alps); <bold>46</bold>, <italic>Capnia nigra</italic> (Greater Geneva region); <bold>47</bold>, <italic>Zwicknia westermanni</italic> (Occitanie); <bold>48</bold>, <italic>Zwicknia bifrons</italic> (Jura Mountains); <bold>49</bold>, <italic>Zwicknia ledoarei</italic> (Jura Mountains); <bold>50</bold>, <italic>Zwicknia gattolliati</italic> (Apennines); <bold>51</bold>, <italic>Zwicknia rupprechti</italic> (Vosges). <bold>52</bold>. Setation on head of <italic>Zwicknia rupprechti</italic> (Vosges).</title>
						<p xml:lang="es">Figs. 45&#x2013;52.&#x2014; <bold>45&#x2013;51</bold>. Vista lateral del epiprocto de larvas macho maduras de <italic>Capnia</italic> y <italic>Zwicknia</italic>: <bold>45</bold>, <italic>Capnia vidua</italic> (Alpes suizos); <bold>46</bold>, <italic>Capnia nigra</italic> (regi&#x00F3;n Gran Ginebra); <bold>47</bold>, <italic>Zwicknia westermanni</italic> (Occitania); <bold>48</bold>, <italic>Zwicknia bifrons</italic> (Monta&#x00F1;as del Jura); <bold>49</bold>, <italic>Zwicknia ledoarei</italic> (Monta&#x00F1;as del Jura); <bold>50</bold>, <italic>Zwicknia gattolliati</italic> (Apeninos); <bold>51</bold>, <italic>Zwicknia rupprechti</italic> (Vosgos). <bold>52</bold>. Setaci&#x00F3;n de la cabeza de <italic>Zwicknia rupprechti</italic> (Vosgos).</p>
					</caption>
					<graphic id="gra-9-e150" xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="GRAELL-77-2-e150-f9-G.png"/>
				</fig>
				<fig id="fig-10-e150">
					<label>Figs. 53&#x2013;55</label>
					<caption>
						<title>Wing venation of <italic>Zwicknia</italic> (annotations by Dr. Olivier B&#x00E9;thoux): <bold>53</bold>, wing venation of micropterous male of <italic>Zwicknia gattolliati</italic>; <bold>54</bold>, wing venation of brachypterous male of <italic>Zwicknia westermanni</italic>; <bold>55</bold>, wing venation of macropterous female of <italic>Zwicknia ledoarei</italic>.</title>
						<p xml:lang="es">Figs. 53&#x2013;55.&#x2014; Venaci&#x00F3;n alar de <italic>Zwicknia</italic> (anotaciones del Dr. Olivier B&#x00E9;thoux): <bold>53</bold>, venaci&#x00F3;n alar del macho micr&#x00F3;ptero de <italic>Zwicknia gattolliati</italic>; <bold>54</bold>, venaci&#x00F3;n alar del macho braqu&#x00ED;ptero de <italic>Zwicknia westermanni</italic>; <bold>55</bold>, venaci&#x00F3;n alar de la hembra macr&#x00F3;ptera de <italic>Zwicknia ledoarei</italic>.</p>
					</caption>
					<graphic id="gra-10-e150" xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="GRAELL-77-2-e150-f10-G.png"/>
				</fig>
				<p>
					<italic>Larvae</italic> (<xref ref-type="fig" rid="fig-4-e150">Figs. 15</xref>&#x2013;<xref ref-type="fig" rid="fig-5-e150">26</xref>). Male and female larvae of <italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold> bear very short and dense erect bristles of nearly equal length on head and pronotum (<xref ref-type="fig" rid="fig-4-e150">Figs. 15&#x2013;17</xref>). Setation on basal segments of antennae very short and sparse. Abdominal tergites with very short, but compact bristles (<xref ref-type="fig" rid="fig-4-e150">Figs. 18&#x2013;19</xref>). Row of bristles on abdominal tergites medially interrupted (<xref ref-type="fig" rid="fig-4-e150">Fig. 18</xref>). Lateral bristles on lower margin of abdominal tergites very short, not reaching the adjacent tergite (<xref ref-type="fig" rid="fig-4-e150">Fig. 19</xref>). Dorsal margin of femur of posterior leg with an uncompacted row of both long and short setae, the longest measuring less than a third of the total width of the femur (<xref ref-type="fig" rid="fig-4-e150">Fig. 20</xref>). Dorsal edge of tibia of hindleg with a narrow row of long and fine setae in its distal part (<xref ref-type="fig" rid="fig-5-e150">Figs. 21&#x2013;22</xref>). Micropterism clearly prefigured on male larvae (<xref ref-type="fig" rid="fig-5-e150">Fig. 23</xref>). In male larvae and nymphs, tergite 10, which prefigures the male epiproct and the process on tergite 9, with wide basis and blunt apex, in dorsal view (<xref ref-type="fig" rid="fig-5-e150">Fig. 24</xref>). In lateral view, tergite 10 is smoothly truncated in near mature larvae (<xref ref-type="fig" rid="fig-5-e150">Fig. 25</xref>) and more pointed in mature larvae (<xref ref-type="fig" rid="fig-5-e150">Fig. 26</xref>).</p>
			</sec>
			<sec id="sec-3.3-e150">
				<title>D<sc>erivatio nominis</sc></title>
				<p>The species is named after the region where it was collected, namely the &#x00C1;vila Province in Spain. The epithet is formed by adding the Latin toponymic suffix &#x2013;<italic>ensis</italic> to the place name &#x00C1;vila and is to be treated as an epicene Latin adjective in the nominative case combined with <italic>Zwicknia</italic>.</p>
			</sec>
			<sec id="sec-3.4-e150">
				<title>M<sc>orphological affinities</sc></title>
				<p>
					<italic>Adult males</italic>. From the morphological point of view, <italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold> stands rather isolated and differs from all other known species or subspecies of the genus <italic>Zwicknia</italic> from Western and Southern Europe by its headward pointing process of tergite 9 in lateral view (<xref ref-type="fig" rid="fig-1-e150">Figs. 1&#x2013;4</xref>), whereas this process is raised perpendicularly in all other known species from these regions (<xref ref-type="fig" rid="fig-6-e150">Figs. 27&#x2013;31</xref>), except for <italic>Zwicknia rupprechti</italic>, where it projects caudally (<xref ref-type="fig" rid="fig-6-e150">Fig. 32</xref>). In lateral view, the process on tergite 9 is highly elevated in <italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold> (<xref ref-type="fig" rid="fig-1-e150">Figs. 1&#x2013;4</xref>), whereas it is low in <italic>Zwicknia rupprechti</italic> (<xref ref-type="fig" rid="fig-6-e150">Fig. 32</xref>) and <italic>Z. ledoarei</italic> (<xref ref-type="fig" rid="fig-6-e150">Fig. 31</xref>). In lateral view, the main epiproct sclerite of <italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold> is slightly pointed (<xref ref-type="fig" rid="fig-1-e150">Figs. 1&#x2013;4</xref>), but not acute as in <italic>Z. acuta</italic> (<xref ref-type="fig" rid="fig-6-e150">Fig. 28</xref>), <italic>Z. westermanni</italic> (<xref ref-type="fig" rid="fig-6-e150">Fig. 29</xref>) and <italic>Zwicknia gattolliati</italic> (<xref ref-type="fig" rid="fig-6-e150">Fig. 30</xref>). The epiproct tip of <italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold> is straight (<xref ref-type="fig" rid="fig-1-e150">Figs 1&#x2013;4</xref>), not upcurved as in <italic>Zwicknia gattolliati</italic> (<xref ref-type="fig" rid="fig-6-e150">Fig. 30</xref>) or <italic>Z. bifrons</italic> (<xref ref-type="fig" rid="fig-6-e150">Fig. 27</xref>). In lateral (<xref ref-type="fig" rid="fig-1-e150">Figs. 1&#x2013;4</xref>) and dorsal (<xref ref-type="fig" rid="fig-1-e150">Figs. 5</xref>, <xref ref-type="fig" rid="fig-2-e150">7&#x2013;8</xref>) views, the main epiproct sclerite of <italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold> is slender and narrow, whereas it is much thicker and larger in <italic>Zwicknia bifrons</italic> (<xref ref-type="fig" rid="fig-6-e150">Fig. 27</xref>; <xref ref-type="bibr" rid="ref-14-e150">Mur&#x00E1;nyi <italic>et al.</italic>, 2014: 41, fig. 82</xref>) and <italic>Z. rupprechti</italic> (<xref ref-type="fig" rid="fig-6-e150">Fig. 32</xref>). In lateral view, the membranous part of the apex of the main epiproct sclerite of <italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold> is very long (<xref ref-type="fig" rid="fig-1-e150">Figs. 3&#x2013;4</xref>), much longer than the one of <italic>Z. bifrons</italic> (<xref ref-type="fig" rid="fig-6-e150">Fig. 27</xref>), <italic>Z. acuta</italic> (<xref ref-type="fig" rid="fig-6-e150">Fig. 28</xref>), <italic>Z. rupprechti</italic> (<xref ref-type="fig" rid="fig-6-e150">Fig. 32</xref>; <xref ref-type="bibr" rid="ref-14-e150">Mur&#x00E1;nyi <italic>et al.</italic> 2014: figs. 106&#x2013;109 and figs. 127&#x2013;147</xref>), <italic>Zwicknia westermanni</italic> (<xref ref-type="fig" rid="fig-6-e150">Fig. 29</xref>), <italic>Z. gattolliati</italic> (<xref ref-type="fig" rid="fig-6-e150">Fig. 30</xref>) and <italic>Z. ledoarei</italic> (<xref ref-type="fig" rid="fig-6-e150">Fig. 31</xref>). In caudal view, the shape of the process on tergite 9 has a wide base, is highly elevated and has a dome-shaped apex (<xref ref-type="fig" rid="fig-1-e150">Fig. 6</xref>), not resembling any of the numerous illustrations of this feature provided by <xref ref-type="bibr" rid="ref-14-e150">Mur&#x00E1;nyi <italic>et al.</italic> (2014: 61, figs. 148&#x2013;164</xref>). The ventral vesicle of <italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold> is very large and rounded (<xref ref-type="fig" rid="fig-2-e150">Fig. 10</xref>), much larger than the one of <italic>Z. ledoarei</italic> (<xref ref-type="fig" rid="fig-7-e150">Fig. 37</xref>; <xref ref-type="bibr" rid="ref-19-e150">Reding <italic>et al.</italic> 2016: 141, fig. 5</xref>) or <italic>Z. rupprechti</italic> (<xref ref-type="fig" rid="fig-7-e150">Fig. 38</xref>; <xref ref-type="bibr" rid="ref-14-e150">Mur&#x00E1;nyi <italic>et al.</italic> 2014: 48, fig. 91</xref>), but not mussel-shaped as the one of <italic>Z. gattolliati</italic> (<xref ref-type="fig" rid="fig-7-e150">Fig. 36</xref>) and <italic>Z. acuta</italic> (<xref ref-type="fig" rid="fig-7-e150">Fig. 34</xref>; <xref ref-type="bibr" rid="ref-14-e150">Mur&#x00E1;nyi <italic>et al.</italic> 2014: 39, fig. 79</xref>), and not elliptical as the one of <italic>Z. bifrons</italic> (<xref ref-type="fig" rid="fig-7-e150">Fig. 33</xref>; <xref ref-type="bibr" rid="ref-14-e150">Mur&#x00E1;nyi <italic>et al.</italic> 2014: 41, fig. 83</xref>) and <italic>Z. westermanni</italic> (<xref ref-type="fig" rid="fig-7-e150">Fig. 35</xref>). Note that the ventral vesicle, that males use for generating their bioacoustic drumming signals, is not visible in part of the specimens, because retracted, partly or entirely, under sternite 8 until the start of the drumming activity. The main epiproct sclerite of <italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold> has a small &#x201C;hunchback&#x201D; in its median dorsal region (<xref ref-type="fig" rid="fig-1-e150">Figs. 3, 4</xref>, arrow), a feature not seen on any other presently described species of <italic>Zwicknia</italic> (<xref ref-type="fig" rid="fig-6-e150">Figs. 27&#x2013;32</xref>) from Western and Southern Europe.</p>
				<p>
					<italic>Adult females</italic>. In <italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold>, the posterior edge of the female subgenital plate is crescent-shaped (<xref ref-type="fig" rid="fig-2-e150">Fig. 12</xref>), whereas it is nearly straight in other species, such as <italic>Z. bifrons</italic> (<xref ref-type="fig" rid="fig-8-e150">Fig. 41</xref>), <italic>Z. ledoarei</italic> (<xref ref-type="fig" rid="fig-8-e150">Fig. 42</xref>), <italic>Z. gattolliati</italic> (<xref ref-type="fig" rid="fig-8-e150">Fig. 43</xref>) and <italic>Z. westermanni</italic> (<xref ref-type="fig" rid="fig-8-e150">Fig. 44</xref>), except for <italic>Z. acuta</italic> (<xref ref-type="bibr" rid="ref-14-e150">Mur&#x00E1;nyi <italic>et al.</italic> 2014: 34, fig. 64</xref>), where it is also crescent-shaped. Like all other known species of <italic>Zwicknia, Z. gattolliati</italic> excepted (<xref ref-type="fig" rid="fig-8-e150">Fig. 40</xref>), female adults of <italic>Z. avilensis</italic>
					<bold>sp. n.</bold> have short bristles on the dorsal edge of femora and tibiae of all three legs (<xref ref-type="fig" rid="fig-2-e150">Fig. 11</xref>; cf. <xref ref-type="fig" rid="fig-8-e150">Fig. 39</xref>). <italic>Zwicknia gattolliati</italic> is, to the present knowledge, the only species of the Western Palearctic whose females have long setae on the dorsal edge of their femora and tibiae (<xref ref-type="fig" rid="fig-8-e150">Fig. 40</xref>; <xref ref-type="bibr" rid="ref-26-e150">Vin&#x00E7;on &#x0026; Reding 2018: figs. 8 and 11</xref>; <xref ref-type="bibr" rid="ref-18-e150">Reding 2021</xref>).</p>
				<p>
					<italic>Larvae</italic>. In larvae of <italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold>, the erect bristles on head and pronotum are dense, compact and very short (<xref ref-type="fig" rid="fig-4-e150">Figs. 15&#x2013;17</xref>), even the shortest among all other larvae of <italic>Zwicknia</italic> examined so far (<italic>Z. bifrons</italic>, <italic>Z. westermanni</italic>, <italic>Z. ledoarei</italic>, <italic>Z. rupprechti</italic> and <italic>Z. gattolliati</italic>; <xref ref-type="bibr" rid="ref-17-e150">Reding 2020: Vol. 4, figs. 17A2, 17A14, 17B9, 17C3 and 17C4</xref>). The bristles on the head of <italic>Zwicknia rupprechti</italic> are also very short, but uncompacted (<xref ref-type="fig" rid="fig-9-e150">Fig. 52</xref>; cf. <xref ref-type="fig" rid="fig-4-e150">Figs. 15, 17</xref>). In lateral view, tergite 10 of mature male larvae of <italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold> is smoothly truncated (<xref ref-type="fig" rid="fig-5-e150">Fig. 25</xref>), or slightly pointed (<xref ref-type="fig" rid="fig-5-e150">Fig. 26</xref>), and different from those of <italic>Capnia vidua</italic> (<xref ref-type="fig" rid="fig-9-e150">Fig. 45</xref>), <italic>Capnia nigra</italic> (<xref ref-type="fig" rid="fig-9-e150">Fig. 46</xref>), <italic>Zwicknia westermanni</italic> (<xref ref-type="fig" rid="fig-9-e150">Fig. 47</xref>), <italic>Z. bifrons</italic> (<xref ref-type="fig" rid="fig-9-e150">Fig. 48</xref>), <italic>Z. ledoarei</italic> (<xref ref-type="fig" rid="fig-9-e150">Fig. 49</xref>), <italic>Z. gattolliati</italic> (<xref ref-type="fig" rid="fig-9-e150">Fig. 50</xref>) and <italic>Z. rupprechti</italic> (<xref ref-type="fig" rid="fig-9-e150">Fig. 51</xref>).</p>
			</sec>
			<sec id="sec-3.5-e150">
				<title>D<sc>istribution and ecology</sc></title>
				<p>The type material of <italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold> has been collected above 1000 m in the Sierra de la Serrota, located within the mountain ranges of the Sistema Central, in small temporary brooks in the headwaters of the Duero basin (<xref ref-type="fig" rid="fig-11-e150">Fig. 56</xref>). The collection dates for the species indicate a winter flight period, from December to March. Isolated females, however, are still found in April, when no males are on the wing anymore. The overall distribution of <italic>Zwicknia avilensis</italic>
					<bold>sp. n.</bold> in Spain is presently unknown. The species inhabits brooks and rivers that dry up in summer (<xref ref-type="fig" rid="fig-11-e150">Fig. 56</xref>; <xref ref-type="bibr" rid="ref-9-e150">Gonz&#x00E1;lez del T&#x00E1;nago 1984</xref>). In December 2020, the species was found with <italic>Leuctra maroccana</italic> Aubert, 1956c and <italic>Capnioneura libera</italic>; in April 2021 with <italic>Brachyptera arcuata</italic> (Klap&#x00E1;lek, 1902) and <italic>Protonemura asturica</italic> (Aubert, 1954).</p>
				<fig id="fig-11-e150">
					<label>Fig. 56</label>
					<caption>
						<title>Collecting locality of <italic>Zwicknia avilensis</italic>
							<bold>sp. n.</bold> (Sierra de la Serrota, Arroyo de Canto Moreno, Mu&#x00F1;otello).</title>
						<p xml:lang="es">Fig. 56.&#x2014; Localidad de recolecci&#x00F3;n de <italic>Zwicknia avilensis</italic>
							<bold>sp. n.</bold> (Sierra de la Serrota, Arroyo de Canto Moreno, Mu&#x00F1;otello).</p>
					</caption>
					<graphic id="gra-11-e150" xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="GRAELL-77-2-e150-f11-G.png"/>
				</fig>
			</sec>
			<sec id="sec-3.6-e150">
				<title>C<sc>onservation status</sc></title>
				<p>In 1981, the species was reported from 14 different sampling stations in the headwaters of river Duero (<xref ref-type="bibr" rid="ref-9-e150">Gonz&#x00E1;lez del T&#x00E1;nago 1984</xref>). In 2020 and 2021, the species is still extant in at least one of its former locations (samplings by the second author). Further collecting is needed to ascertain the present conservation status of the species.</p>
				<p>
					<bold>Identification key for adult <italic>Zwicknia</italic> from Western and Southern Europe</bold>
				</p>
				<list list-type="simple">
					<list-item>
						<p>1 Characteristic process on tergite 9 (<xref ref-type="fig" rid="fig-1-e150">Figs. 1&#x2013;4, 5</xref>, <xref ref-type="fig" rid="fig-6-e150">27&#x2013;32</xref>). Presence of a ventral vesicle at the center of the posterior margin of sternite 8 (<xref ref-type="fig" rid="fig-2-e150">Figs. 10</xref>, <xref ref-type="fig" rid="fig-7-e150">33&#x2013;38</xref>). Micropterous (<xref ref-type="fig" rid="fig-2-e150">Figs. 9</xref>, <xref ref-type="fig" rid="fig-10-e150">53</xref>) or brachypterous (<xref ref-type="fig" rid="fig-10-e150">Fig. 54</xref>) ___ <bold>3 (male adults of <italic>Zwicknia</italic>)</bold>
						</p>
					</list-item>
					<list-item>
						<p>&#x2013; No characteristic process on tergite 9. Macropterous (or slightly brachypterous in some specimens from the British-Irish Isles), with anterior wing with a reduced number of cross-veins (three) between CuA and M (<xref ref-type="fig" rid="fig-10-e150">Fig. 55</xref>; cf. <xref ref-type="fig" rid="fig-3-e150">Fig. 14</xref>). First cell between CuA and CuP in anterior wings quadrangular in shape (<xref ref-type="fig" rid="fig-3-e150">Figs. 13</xref>, <xref ref-type="fig" rid="fig-10-e150">55</xref>). Posterior margin of subgenital plate (sternite 8) rectilinear, slightly rounded or crescent-shaped (<xref ref-type="fig" rid="fig-2-e150">Figs. 12</xref>, <xref ref-type="fig" rid="fig-8-e150">41&#x2013;44</xref>) ___ <bold>2 (female adults of <italic>Zwicknia</italic>)</bold>
						</p>
					</list-item>
					<list-item>
						<p>2 Femora and tibiae with long and dense setae (<xref ref-type="fig" rid="fig-8-e150">Figs. 40</xref>). Margins of tergites and sternites with a dense row of setae, well visible in ventral view (<xref ref-type="fig" rid="fig-8-e150">Fig. 43</xref>) ___ <bold>
								<italic>Zwicknia gattolliati</italic>
							</bold>
						</p>
					</list-item>
					<list-item>
						<p>&#x2013; Femora and tibiae with only short setae (<xref ref-type="fig" rid="fig-2-e150">Figs. 11</xref>, <xref ref-type="fig" rid="fig-8-e150">39</xref>). Margins of tergites and sternites without a dense row of setae well visible in ventral view (<xref ref-type="fig" rid="fig-2-e150">Figs. 12</xref>, <xref ref-type="fig" rid="fig-8-e150">41&#x2013;42, 44</xref>) ___ <bold>other female adults of <italic>Zwicknia</italic> (not keyed out)</bold>
						</p>
					</list-item>
					<list-item>
						<p>3 Brachypterous (<xref ref-type="fig" rid="fig-10-e150">Fig. 54</xref>). Veins R and M not fused in the proximal part of the forewings (<xref ref-type="fig" rid="fig-10-e150">Fig. 54</xref>) ___ <bold>4</bold>
						</p>
					</list-item>
					<list-item>
						<p>&#x2013; Micropterous (<xref ref-type="fig" rid="fig-2-e150">Figs. 9</xref>, <xref ref-type="fig" rid="fig-10-e150">53</xref>). Veins R and M (and sometimes also ScP) fused at least in the proximal part of the forewings (<xref ref-type="fig" rid="fig-10-e150">Fig. 53</xref>)  ___ <bold>5</bold>
						</p>
					</list-item>
					<list-item>
						<p>4 Main epiproct sclerite very long and slender, almost reaching tergite 7 (<xref ref-type="fig" rid="fig-6-e150">Fig. 29</xref>). Tip of main epiproct sclerite straight, not upcurved, long, narrow, regularly bent and pointed at apex, in lateral view (<xref ref-type="fig" rid="fig-6-e150">Fig. 29</xref>). Process on tergite 9 highly elevated and perpendicularly raised (<xref ref-type="fig" rid="fig-6-e150">Fig. 29</xref>). Ventral vesicle large, half as wide as the subgenital plate and elliptical (<xref ref-type="fig" rid="fig-7-e150">Fig. 35</xref>) ___ <bold>
								<italic>Zwicknia westermanni</italic>
							</bold>
						</p>
					</list-item>
					<list-item>
						<p>5 Process of tergite 9 projecting caudally (<xref ref-type="fig" rid="fig-6-e150">Fig. 32</xref>) or orientated headward (<xref ref-type="fig" rid="fig-1-e150">Figs. 1&#x2013;4</xref>)  ___ <bold>6</bold>
						</p>
					</list-item>
					<list-item>
						<p>&#x2013; Process of tergite 9 perpendicularly elevated, not projecting caudally or headward (<xref ref-type="fig" rid="fig-6-e150">Figs. 27&#x2013;31</xref>) ___ <bold>7</bold>
						</p>
					</list-item>
					<list-item>
						<p>6 Process of tergite 9 projecting caudally (<xref ref-type="fig" rid="fig-6-e150">Fig. 32</xref>). Tip of main epiproct sclerite large and pointed in lateral view (<xref ref-type="fig" rid="fig-6-e150">Fig. 32</xref>). Ventral vesicle small and rounded (<xref ref-type="fig" rid="fig-7-e150">Fig. 38</xref>)___ <bold>
								<italic>Zwicknia rupprechti</italic>
							</bold>
						</p>
					</list-item>
					<list-item>
						<p>&#x2013; Process of tergite 9 projecting headward (<xref ref-type="fig" rid="fig-1-e150">Figs. 1&#x2013;4</xref>). Tip of main epiproct sclerite straight, neither downcurved nor upcurved and blunt (<xref ref-type="fig" rid="fig-1-e150">Figs. 1&#x2013;4</xref>). Ventral vesicle very large and rounded (<xref ref-type="fig" rid="fig-2-e150">Fig. 10</xref>) ___ <bold>
								<italic>Zwicknia avilensis</italic> sp. n.</bold>
						</p>
					</list-item>
					<list-item>
						<p>7 Process of tergite 9 very low, its height not reaching half the length of the corresponding tergite (<xref ref-type="fig" rid="fig-6-e150">Fig. 31</xref>). In lateral view, the tip of the main epiproct sclerite is blunt and nearly rounded (<xref ref-type="fig" rid="fig-6-e150">Fig. 31</xref>). Ventral vesicle very small and rounded (<xref ref-type="fig" rid="fig-7-e150">Fig. 37</xref>) ___ <bold>
								<italic>Zwicknia ledoarei</italic>
							</bold>
						</p>
					</list-item>
					<list-item>
						<p>&#x2013; Process of tergite 9 high, its height higher than half the length of the corresponding tergite 9 (<xref ref-type="fig" rid="fig-6-e150">Figs. 27&#x2013;28, 30</xref>) ___ <bold>8</bold>
						</p>
					</list-item>
					<list-item>
						<p>8 In lateral view, the tip of the main epiproct sclerite is slender, long, narrow and pointed (<xref ref-type="fig" rid="fig-6-e150">Figs. 28, 30</xref>) ___ <bold>9</bold>
						</p>
					</list-item>
					<list-item>
						<p>&#x2013; In lateral view, the tip of the main epiproct sclerite is bulky, pointed, wide-angled and short, slightly up-curved (<xref ref-type="fig" rid="fig-6-e150">Fig. 27</xref>). Process of tergite 9 very high and perpendicularly elevated (<xref ref-type="fig" rid="fig-6-e150">Fig. 27</xref>). Ventral vesicle large and elliptical in shape (<xref ref-type="fig" rid="fig-7-e150">Fig. 33</xref>) ___ <bold>
								<italic>Zwicknia bifrons</italic>
							</bold>
						</p>
					</list-item>
					<list-item>
						<p>9 The tip of the main epiproct sclerite is straight, not up-curved (<xref ref-type="fig" rid="fig-6-e150">Fig. 28</xref>). Main epiproct sclerite very slender and thin (<xref ref-type="fig" rid="fig-6-e150">Fig. 28</xref>). Ventral vesicle large and mussel-shaped (<xref ref-type="fig" rid="fig-7-e150">Fig. 34</xref>) ___ <bold>
								<italic>Zwicknia acuta</italic>
							</bold>
						</p>
					</list-item>
					<list-item>
						<p>&#x2013; The tip of the main epiproct sclerite is very slightly up-curved (<xref ref-type="fig" rid="fig-6-e150">Fig. 30</xref>). Ventral vesicle medium-sized and mussel-shaped (<xref ref-type="fig" rid="fig-7-e150">Fig. 36</xref>) ___ <bold>
								<italic>Zwicknia gattolliati</italic>
							</bold>
						</p>
					</list-item>
				</list>
			</sec>
		</sec>
		<sec sec-type="discussion" id="sec-4-e150">
			<title>Discussion</title>
			<p>
				<xref ref-type="bibr" rid="ref-15-e150">Nav&#x00E1;s (1917)</xref> mentions two new species of <italic>Capnia</italic> from Spain, <italic>C. dusmeti</italic> and <italic>C. maynari</italic>. The corresponding type specimens are lost (<xref ref-type="bibr" rid="ref-2-e150">Aubert 1956a</xref>). According to <xref ref-type="bibr" rid="ref-3-e150">Aubert (1956b)</xref>, <italic>Capnia maynari</italic> is a synonym of <italic>C. nigra</italic>, and <italic>Capnia dusmeti</italic> a synonym of <italic>Capnia bifrons</italic>. <italic>Capnia maynari</italic> had been described by <xref ref-type="bibr" rid="ref-15-e150">Nav&#x00E1;s (1917)</xref> on the basis of a single female, but the drawing of the wing venation in <xref ref-type="bibr" rid="ref-15-e150">Nav&#x00E1;s (1917: fig. 4</xref>; cf. <xref ref-type="fig" rid="fig-2-e150">Fig. 14</xref>) clearly shows that the first cell between CuA and CuP in the anterior wing is triangular in shape. <italic>Capnia maynari</italic> hence belongs to the genus <italic>Capnia</italic>, while <italic>C. dusmeti</italic>, also described by <xref ref-type="bibr" rid="ref-15-e150">Nav&#x00E1;s (1917)</xref> from a single female, belongs to the genus <italic>Zwicknia</italic>, since the first cell between CuA and CuP in the anterior wing of this specimen is quadrangular in shape (cf. <xref ref-type="fig" rid="fig-3-e150">Figs. 13</xref>, <xref ref-type="fig" rid="fig-10-e150">55</xref>; <xref ref-type="bibr" rid="ref-15-e150">Nav&#x00E1;s 1917: fig. 3</xref>; <xref ref-type="bibr" rid="ref-4-e150">Aubert 1959: 26, fig. 22</xref>; <xref ref-type="bibr" rid="ref-17-e150">Reding 2020: vol. I V, fig. 17A20</xref>). <italic>Capnia</italic> (<italic>Zwicknia</italic>) <italic>dusmeti</italic> had been collected at the Escorial, Sierra de Guadarrama, on March 26<sup>th</sup> 1916. The taxonomic identity of <italic>Capnia dusmeti</italic> has never been clearly established and the taxon is considered as a <italic>nomen dubium</italic> by <xref ref-type="bibr" rid="ref-14-e150">Mur&#x00E1;nyi <italic>et al.</italic> (2014: 56)</xref>, as a synonym of <italic>Capnia bifrons</italic> by <xref ref-type="bibr" rid="ref-1-e150">Aubert (1952: 247</xref>; <xref ref-type="bibr" rid="ref-3-e150">1956b</xref>) and <xref ref-type="bibr" rid="ref-11-e150">Illies (1966)</xref>, and as a synonym of <italic>Zwicknia bifrons</italic> by <xref ref-type="bibr" rid="ref-8-e150">DeWalt <italic>et al.</italic> (2021)</xref>. In the past, the now obsolete taxon <italic>Capnia bifrons</italic> has been mentioned from Spain by many authors (<xref ref-type="bibr" rid="ref-15-e150">Nav&#x00E1;s 1917</xref>; <xref ref-type="bibr" rid="ref-16-e150">Puig Garc&#x00ED;a 1984</xref>; <xref ref-type="bibr" rid="ref-9-e150">Gonz&#x00E1;lez del T&#x00E1;nago 1984</xref>; <xref ref-type="bibr" rid="ref-21-e150">S&#x00E1;nchez-Ortega &#x0026; Tierno de Figueroa 1996</xref>; <xref ref-type="bibr" rid="ref-12-e150">Luz&#x00F3;n-Ortega &#x0026; Tierno de Figueroa 2000</xref>; <xref ref-type="bibr" rid="ref-22-e150">Tierno de Figueroa <italic>et al.</italic> 2003</xref>; <xref ref-type="bibr" rid="ref-25-e150">Vin&#x00E7;on &#x0026; Pardo 2004</xref>; <xref ref-type="bibr" rid="ref-10-e150">Graf <italic>et al.</italic> 2009</xref>; <xref ref-type="bibr" rid="ref-13-e150">Luz&#x00F3;n-Ortega <italic>et al</italic>. 2013</xref>). Since these records have not yet been revised after the breaking up of the former taxon <italic>Capnia bifrons</italic> into different species of the genus <italic>Zwicknia</italic>, it is presently not possible to decide whether all of them belong to <italic>Zwicknia avilensis</italic>
				<bold>sp. n.</bold>, or to different, currently described or yet unknown species, of the genus <italic>Zwicknia</italic> (<xref ref-type="bibr" rid="ref-23-e150">Tierno de Figueroa <italic>et al</italic>. 2014</xref>, <xref ref-type="bibr" rid="ref-24-e150">2018</xref>). Previous hints as to the possible existence of unknown species of <italic>Zwicknia</italic> in Spain had been provided by <xref ref-type="bibr" rid="ref-20-e150">Rupprecht (1997)</xref>, <xref ref-type="bibr" rid="ref-14-e150">Mur&#x00E1;nyi <italic>et al</italic>. (2014: 56)</xref> and <xref ref-type="bibr" rid="ref-7-e150">Boumans &#x0026; Mur&#x00E1;nyi (2014)</xref>, on the basis of drumming and genetic studies.</p>
		</sec>
	</body>
	<back>
		<ack id="ack-1-e150">
			<title>Acknowledgements</title>
			<p>We are grateful to Dr. D&#x00E1;vid Mur&#x00E1;nyi (Department of Zoology, Institute of Biology, Eszterh&#x00E1;zy K&#x00E1;roly University, Le&#x00E1;nyka &#x00FA;t 6, H-3300 Eger, Hungary) for helpful and constructive comments as well as for providing paratypes of <italic>Zwicknia acuta</italic> from Slovakia. We are furthermore indebted to Dr. Jos&#x00E9; Manuel Tierno de Figueroa (Department of Zoology, University of Granada, Spain) for the loan of specimens of <italic>Zwicknia avilensis</italic>
				<bold>sp. n.</bold>, from the S&#x00E1;nchez-Ortega collection and for the revision of the manuscript. Special thanks also to Dr. Olivier B&#x00E9;thoux (Centre de Recherche sur la Pal&#x00E9;obiodiversit&#x00E9; et les Pal&#x00E9;oenvironnements (CR2P), Mus&#x00E9;um National dHistoire Naturelle, Paris, France) for his help in annotating the wing venation of capniid stoneflies. Figures 35, 37, 42, 45, 46 and 47 are modified from those produced by the first author from <italic>Zootaxa</italic> 4382(2) and figures 36, 39, 43 and 53 from <italic>Zootaxa</italic> 4486(1) with permission of Magnolia Press.</p>
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