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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.v77.319</article-id>
			<article-id pub-id-type="publisher-id">graellsia.2021.v77.319</article-id>
			<article-categories>
				<subj-group subj-group-type="heading">
					<subject>Notas / <italic>Notes</italic></subject>
				</subj-group>
			</article-categories>
			<title-group>
				<article-title>On the larva of the Iberian allochthonous endemic firefly <italic>Photinus immigrans</italic>
					Zaragoza-Caballero &#x0026; Vi&#x00F1;olas, 2018 (Coleoptera, Lampyridae)</article-title>
				<trans-title-group xml:lang="es">
					<trans-title>Sobre la larva de la luci&#x00E9;rnaga ib&#x00E9;rica end&#x00E9;mica <italic>Photinus immigrans</italic>
						Zaragoza-Caballero &#x0026; Vi&#x00F1;olas, 2018 (Coleoptera, Lampyridae)</trans-title>
				</trans-title-group>
				<alt-title alt-title-type="running-head">Notas / <italic>Notes</italic></alt-title>
			</title-group>
			<contrib-group>
				<contrib contrib-type="author" corresp="yes">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0003-1555-9598</contrib-id>
					<name name-style="western">
						<surname>Rodr&#x00ED;guez-Flores</surname>
						<given-names>Paula C.</given-names>
					</name>
					<xref ref-type="aff" rid="aff-1-e154">1</xref>
					<xref ref-type="aff" rid="aff-2-e154">2</xref>
					<aff id="aff-1-e154">
						<label>1</label>
						<institution>Department of Organismic and Evolutionary Biology, Museum of Comparative Zoology, Harvard University</institution>, <addr-line>Cambridge, MA</addr-line>, <country>USA</country>
					</aff>
					<aff id="aff-2-e154">
						<label>2</label>
						<institution>Centre d'Estudis Avan&#x00E7;ats de Blanes (CEAB-CSIC)</institution>, <addr-line>C. acc. Cala Sant Francesc 14 17300 Blanes, Girona</addr-line>, <country>Spain</country>
					</aff>
					<email xlink:href="paularodriguezflores@gmail.com">paularodriguezflores@gmail.com</email>
				</contrib>
				<contrib contrib-type="author">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0001-8086-8667</contrib-id>
					<name name-style="western">
						<surname>Recuero</surname>
						<given-names>Ernesto</given-names>
					</name>
					<xref ref-type="aff" rid="aff-3-e154">3</xref>
					<aff id="aff-3-e154">
						<label>3</label>
						<institution>Department of Plant &#x0026; Environmental Sciences, Clemson University</institution>, <addr-line>Clemson, SC 29634</addr-line>, <country>USA</country>
					</aff>
					<email xlink:href="ernestorecuerogil@gmail.com">ernestorecuerogil@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>e154</elocation-id>
			<history>
				<date date-type="received">
					<day>07</day>
					<month>06</month>
					<year>2021</year>
				</date>
				<date date-type="accepted">
					<day>27</day>
					<month>10</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-e154.pdf"/>
			<abstract>
				<p>The firefly species <italic>Photinus immigrans</italic> is currently known only from the Iberian Peninsula, although it is likely an introduced species that originated from somewhere in North America. Its last larval stage is here newly recorded and illustrated. The larvae can be easily distinguished from the larvae of Iberian native species by having several longitudinal light brown to reddish stripes on each segment on a black to dark brown background. They were found living in the soil, in the upper 10&#x2013;15 cm. <italic>P. immigrans</italic> lives in sympatry with at least three other native lampyrid species: <italic>Lampyris iberica</italic>, <italic>Nyctophila reichii</italic> and <italic>Lamprohiza mulsantii,</italic> although it was not observed in strict syntopy, at least during larval phases, in our surveys. Larvae of <italic>Photinus immigrans</italic> probably use at least partly different alimentary resources, and adult communication signals are different to those of native species living in sympatry, although a detailed study of interspecific competition and the potential effect of the species on the native fireflies is pending.</p>
			</abstract>
			<trans-abstract xml:lang="es">
				<p>La luci&#x00E9;rnaga <italic>Photinus immigrans</italic> se conoce &#x00FA;nicamente en la pen&#x00ED;nsula ib&#x00E9;rica, aunque probablemente es una especie originaria de Norteam&#x00E9;rica. En esta nota presentamos el primer registro e ilustraci&#x00F3;n de su larva. &#x00C9;sta puede ser f&#x00E1;cilmente identificada gracias a su patr&#x00F3;n de coloraci&#x00F3;n, formado en cada segmento por varias l&#x00ED;neas longitudinales marr&#x00F3;n claro o rojizas sobre fondo negro o marr&#x00F3;n oscuro. Se han encontrado viviendo enterradas en capas superficiales del suelo, hasta 10&#x2013;15 cm de profundidad. <italic>P</italic>. <italic>immigrans</italic> vive en simpatr&#x00ED;a con al menos otras tres especies de luci&#x00E9;rnaga nativas de la pen&#x00ED;nsula ib&#x00E9;rica: <italic>Lampyris iberica</italic>, <italic>Nyctophila reichii</italic> and <italic>Lamprohiza mulsantii</italic>, aunque aparentemente no hay sintop&#x00ED;a estricta al menos durante las fases larvarias. Las larvas de <italic>P. immigrans</italic> probablemente utilizan recursos tr&#x00F3;ficos parcialmente diferentes a los de las especies nativas, y su se&#x00F1;alizaci&#x00F3;n luminosa es tambi&#x00E9;n muy diferente a aquellas de otras especies simp&#x00E1;tricas, aunque es necesario un estudio en profundidad sobre competencia interespec&#x00ED;fica y el posible efecto de esta especie ex&#x00F3;tica sobre las luci&#x00E9;rnagas nativas.</p>
			</trans-abstract>
			<kwd-group>
				<kwd>alien species</kwd>
				<kwd>identification</kwd>
				<kwd>sympatry</kwd>
				<kwd>interspecific competition</kwd>
				<kwd>larval stage</kwd>
			</kwd-group>
			<kwd-group xml:lang="es">
				<kwd>especie introducida</kwd>
				<kwd>identificaci&#x00F3;n</kwd>
				<kwd>simpatr&#x00ED;a</kwd>
				<kwd>competencia interespec&#x00ED;fica</kwd>
				<kwd>estado larvario</kwd>
			</kwd-group>
			<counts>
				<page-count count="4"/>
			</counts>
		</article-meta>
	</front>
	<body>
		<p>Fireflies of the American genus <italic>Photinus</italic> Laporte, 1833 are widely distributed through North America, and particularly diverse in Mexico (<xref ref-type="bibr" rid="ref-7-e154">Green, 1956</xref>; <xref ref-type="bibr" rid="ref-15-e154">Zaragoza-Caballero, 1996</xref>; <xref ref-type="bibr" rid="ref-16-e154">Zaragoza-Caballero &#x0026; Vi&#x00F1;olas, 2018</xref>; <xref ref-type="bibr" rid="ref-17-e154">Zaragoza-Caballero <italic>et al</italic>., 2020</xref>). Recently, a new species of <italic>Photinus</italic> was described from the Iberian Peninsula, representing the first record of the genus for the European continent (<xref ref-type="bibr" rid="ref-14-e154">Vi&#x00F1;olas <italic>et al</italic>., 2018</xref>; <xref ref-type="bibr" rid="ref-16-e154">Zaragoza-Caballero &#x0026; Vi&#x00F1;olas, 2018</xref>). The species is considered an alien in Europe that most likely comes from Mexico or the USA. It has been recorded in several localities throughout Girona province, Northeastern Iberian Peninsula (<xref ref-type="bibr" rid="ref-16-e154">ZaragozaCaballero &#x0026; Vi&#x00F1;olas, 2018</xref>), where its range might be expanding. Here we present the first record of the larvae of <italic>P. immigrans</italic> and some notes on their natural history.</p>
		<p>During a summer field trip hundreds of adult male <italic>P. immigrans</italic> (<xref ref-type="fig" rid="fig-1-e154">Fig. 1a</xref>) were observed in the locality of Fonolleres, Girona (42.0201&#x00B0;N, 3.0542&#x00B0;E, 14m, August 19<sup>th</sup>, 2020), flying and flashing over corn fields just by the village soon after dark, in proximity with female specimens of the native <italic>Lampyris iberica</italic>
			<xref ref-type="bibr" rid="ref-6-e154">Geisthardt, Figueira, Day &#x0026; De Cock, 2008</xref> (<xref ref-type="fig" rid="fig-1-e154">Fig. 1e</xref>). We visited the same locality in autumn (November 16<sup>th</sup>, 2020) trying to locate and photograph <italic>P</italic>. <italic>immigrans</italic> larvae, but we could only find larvae of <italic>Lamprohiza mulsantii</italic> (Kiesenwetter, 1850) (<xref ref-type="fig" rid="fig-1-e154">Fig. 1c</xref>) and <italic>Lampyris iberica</italic>. In the nearby locality of Ullastret, Girona (42.0089&#x00B0;N, 3.0778&#x00B0;E, 20m), in an area with wheat crops and small patches of natural vegetation dominated by <italic>Quercus ilex</italic> L., we also located larvae of <italic>L</italic>. <italic>mulsantii</italic> and <italic>Nyctophila reichii</italic> (Jacquelin Du Val, 1859) (<xref ref-type="fig" rid="fig-1-e154">Fig. 1d</xref>). In a new survey, during late winter at the same spot in Fonolleres (February 24<sup>th</sup>, 2021), we found 11 larvae of <italic>P. immigrans</italic> (<xref ref-type="fig" rid="fig-1-e154">Fig. 1b</xref>), superficially buried in a recently ploughed (less than a week) corn field.</p>
		<fig id="fig-1-e154">
			<label>Fig. 1</label>
			<caption>
				<title>Habitus <italic>in vivo</italic>: a) adult male of <italic>Photinus immigrans</italic> from Fonolleres, Girona; b) larva of <italic>Photinus immigrans</italic> from Fonolleres, Girona; c) larva of <italic>Lamprohiza mulsantii</italic> from Fonolleres, Girona; d) larva of <italic>Nyctophila reichii</italic> from Ullastret, Girona; e) Adult female of <italic>Lampyris iberica</italic> from Fonolleres, Girona; f) Larva of <italic>Lampyris iberica</italic> from Blanes, Girona. Photo credits: E. Recuero (a, e); Paula C. Rodr&#x00ED;guez-Flores (b, c, d, f).</title>
				<p xml:lang="es">Fig. 1.&#x2014; Habitus in vivo: a) macho adulto de <italic>Photinus immigrans</italic> de Fonolleres, Girona; b) larva de <italic>Photinus immigrans</italic> de Fonolleres, Girona; c) larva de <italic>Lamprohiza mulsantii</italic> de Fonolleres, Girona; d) larva de <italic>Nyctophila reichii</italic> de Ullastret, Girona; e) Hembra adulta de <italic>Lampyris iberica</italic> de Fonolleres, Girona; f) Larva de <italic>Lampyris iberica</italic> de Blanes, Girona. Cr&#x00E9;ditos de las fotos: E. Recuero (a, e); Paula C. Rodr&#x00ED;guez-Flores (b, c, d, f).</p>
			</caption>
			<graphic id="gra-1-e154" xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="GRAELL-77-2-e154-f1-G.png"/>
		</fig>
		<p>
			<italic>De visu</italic> description: the larval sizes were ca. 10&#x2013;15 mm length, with elongate vermiform body (<xref ref-type="fig" rid="fig-1-e154">Fig. 1a</xref>); observed larvae had 3 thoracic segments and 8 abdominal segments, all tergites were divided by a sagittal line in dorsal view; coloration was dorsally black to dark brown, each tergite had 2 longitudinal light-brown to reddish stripes laterally oriented and widened at posterior part. All specimens had pink lateral pleura and were ventrally lighter than dorsally. The specimens did not show bioluminescence at any point, when found or when disturbed.</p>
		<p>The larvae of this species were found buried in the removed soil of a ploughed field, in the first 10&#x2013;15 cm. By comparison, the native firefly larvae were found in all cases under rocks and debris, and they were not observed in strict syntopy during our surveys. According to the bibliographic records, other species of <italic>Photinus</italic> overwinter for 1&#x2013;2 years as subterranean larvae, and feed mainly on earthworms and soft-bodied insects before pupating during early summer (<xref ref-type="bibr" rid="ref-13-e154">Williams, 1917</xref>; <xref ref-type="bibr" rid="ref-9-e154">Hess, 1920</xref>). This different trophic niche exploitation may account for low trophic competition between <italic>P. immigrans</italic> and the native species. <xref ref-type="bibr" rid="ref-4-e154">De Cock &#x0026; Guzm&#x00E1;n-&#x00C1;lvarez (2013)</xref> reported that some Iberian fireflies occasionally might feed on earthworms, but they preferentially feed on snails and slugs (<xref ref-type="bibr" rid="ref-8-e154">Haddon, 1915</xref>; <xref ref-type="bibr" rid="ref-12-e154">Symondson, 2004</xref>; <xref ref-type="bibr" rid="ref-4-e154">De Cock &#x0026; Guzm&#x00E1;n-&#x00C1;lvarez, 2013</xref>). Mating behavior is also quite dissimilar among <italic>P . immigrans</italic> and most native Iberian species. In <italic>Photinus</italic> fireflies, males produce courtship flashes while flying and females reply to the preferred male with their own flashes (<xref ref-type="bibr" rid="ref-2-e154">Cratsley &#x0026; Lewis, 2003</xref>), whereas males of sympatric native species do not flash during flight and they are usually found flying looking for the continuous cold light produced by apterous females (<xref ref-type="bibr" rid="ref-6-e154">Geisthardt <italic>et al</italic>., 2008</xref>; <xref ref-type="bibr" rid="ref-3-e154">De Cock, 2009</xref>; <xref ref-type="bibr" rid="ref-10-e154">Nov&#x00E1;k, 2017</xref>, <xref ref-type="bibr" rid="ref-11-e154">2018</xref>). We hypothesize that interaction should be low between the alien <italic>P</italic>. <italic>immigrans</italic> and the sympatric native species. However, the species might potentially interfere with species with comparable mating behavior, like <italic>Luciola lusitanica</italic> (Charpentier, 1825), which was recorded historically in the Pyrenees, although its presence is not confirmed in this area (<xref ref-type="bibr" rid="ref-5-e154">De la Fuente, 1931</xref>; <xref ref-type="bibr" rid="ref-4-e154">De Cock &#x0026; Guzm&#x00E1;n-&#x00C1;lvarez, 2013</xref>; <xref ref-type="bibr" rid="ref-1-e154">Constantin, 2014</xref>). A detailed study of interspecific competition and the putative effect of <italic>P</italic>. <italic>immigrans</italic> on the native firefly species and their prey is pending.</p>
		<p>With this contribution we hope to facilitate the identification in the field of larval stages of this exotic species. A detailed description of all stages and larval instars is still needed, for which it is necessary to have access to abundant material including all larval stages. However, the information and images provided in this article will help to identify new populations of <italic>P. immigrans</italic> and track the colonization of new geographic areas during all seasons, as adults are only active in the summer months.</p>
	</body>
	<back>
		<ack id="ack-1-e154">
			<title>Acknowledgements</title>
			<p>Thanks to Miriam Casal and Diego Gil for guiding us in our first visit to the sampling locality, to Michael Caterino for his helpful comments and corrections, and to the two anonymous reviewers that evaluated the manuscript and provided further suggestions and corrections.</p>
		</ack>
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