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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">GRA</journal-id>
<journal-title-group>
<journal-title>Graellsia</journal-title>
</journal-title-group>
<issn pub-type="ppub">0367-5041</issn>
<issn pub-type="epub">1989-953X</issn>
<publisher>
<publisher-name>Consejo Superior de Investigaciones Cient&#x00ED;ficas; Sociedad de Amigos del Museo Nacional de Ciencias Naturales</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="publisher-id">GRA2020_e113</article-id>
<article-id pub-id-type="doi">10.3989/graellsia.2020.v76.267</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Notas / <italic>Notes</italic></subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Range extension of <italic>Harpactirella insidiosa</italic> (Denis, 1960) (Araneae: Mygalomorphae: Theraphosidae): a poorly known endemic spider from Morocco</article-title>
<trans-title-group xml:lang="es">
<trans-title>Ampliaci&#x00F3;n del rango de distribuci&#x00F3;n de <italic>Harpactirella insidiosa</italic> (Denis, 1960) (Araneae: Mygalomorphae: Theraphosidae): una ara&#x00F1;a end&#x00E9;mica de Marruecos poco conocida</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">
<name>
<surname>Calatayud-Mascarell</surname>
<given-names>Arnau</given-names>
</name>
<xref ref-type="aff" rid="aff0001">1</xref>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>S&#x00E1;nchez-Vialas</surname>
<given-names>Alberto</given-names>
</name>
<xref ref-type="aff" rid="aff0001">1</xref>
<xref ref-type="corresp" rid="cor1">&#x002A;</xref>
</contrib>
</contrib-group>
<aff id="aff0001"><label>1</label><institution>Museo Nacional de Ciencias Naturales, MNCN-CSIC</institution>, <addr-line>c/Jos&#x00E9; Guti&#x00E9;rrez Abascal, 2, 28006, Madrid</addr-line>, <country>Spain</country></aff>
<author-notes>
<corresp id="cor1"><label>&#x002A;</label>Corresponding author: <email xlink:href="alberto.alytes@gmail.com">alberto.alytes@gmail.com</email></corresp>
<fn><p>ORCID iD: AC-M: <ext-link ext-link-type="uri" xlink:href="http://orcid.org/0000-0003-0627-2616">http://orcid.org/0000-0003-0627-2616</ext-link> &#x2014; AS-V: <ext-link ext-link-type="uri" xlink:href="http://orcid.org/0000-0003-0068-7669">http://orcid.org/0000-0003-0068-7669</ext-link></p></fn>
</author-notes>
<pub-date pub-type="epub">
<day>31</day>
<month>12</month>
<year>2020</year>
</pub-date>
<pub-date pub-type="collection">
<month>12</month>
<year>2020</year>
</pub-date>
<volume>76</volume>
<issue>2</issue>
<elocation-id content-type="identifier">e113</elocation-id>
<history>
<date date-type="received">
<day>24</day>
<month>01</month>
<year>2020</year>
</date>
<date date-type="accepted">
<day>01</day>
<month>06</month>
<year>2020</year>
</date>
<date date-type="Published online">
<day>09</day>
<month>10</month>
<year>2020</year>
</date>
</history>
<permissions>
<copyright-statement>&#x00A9; 2020 SAM &#x0026; CSIC</copyright-statement>
<copyright-year>2020</copyright-year>
<license license-type="open-access" xlink:href="http://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>
<abstract>
<p>We provide a new record of the poorly studied theraphosid <italic>Harpactirella insidiosa</italic> (Denis, <xref ref-type="bibr" rid="cit0004">1960</xref>) from Morocco, which was previously known only from its type locality in the surroundings of Ben Slimane. The new record, based on two adult females, extends the species distribution 210 km southwest from the type locality. These specimens were found at the Doukkala plain, 14 km southwest of Oualidia, and compared with topotypic specimens. In addition, we provide the first pictures of living specimens of the species and discuss its taxonomic status.</p>
</abstract>
<trans-abstract xml:lang="es">
<p>En este trabajo presentamos una nueva cita de <italic>Harpactirella insidiosa</italic> (Denis, <xref ref-type="bibr" rid="cit0004">1960</xref>), un teraf&#x00F3;sido escasamente estudiado de Marruecos, solamente conocido de la localidad tipo, situada en los alrededores de Ben Slimane. Este nuevo registro, basado en dos hembras adultas, ampl&#x00ED;a el &#x00E1;rea de distribuci&#x00F3;n de la especie 210 km al suroeste de la localidad tipo. Ambos ejemplares fueron encontrados en la llanura de Doukkala, 14 km al sur de Oualidia, y comparados con topotipos de la especie. Esta nota se acompa&#x00F1;a con las primeras im&#x00E1;genes de la especie <italic>in vivo</italic> y se discute, adem&#x00E1;s, su situaci&#x00F3;n taxon&#x00F3;mica.</p>
</trans-abstract>
<kwd-group xml:lang="en">
<kwd><italic>Luphocemus</italic></kwd>
<kwd>spiders</kwd>
<kwd>North Africa</kwd>
<kwd>Doukkala plain</kwd>
<kwd>new record</kwd>
<kwd>geographic distribution</kwd>
</kwd-group>
<kwd-group xml:lang="es">
<kwd><italic>Luphocemus</italic></kwd>
<kwd>ara&#x00F1;as</kwd>
<kwd>Norte de &#x00C1;frica</kwd>
<kwd>llanura de Doukkala</kwd>
<kwd>nuevo registro</kwd>
<kwd>distribuci&#x00F3;n geogr&#x00E1;fica</kwd>
</kwd-group>
</article-meta>
</front>
<body>
<p>The spiders of the family Theraphosidae from Morocco are largely understudied and comprise two genera: <italic>Ischnocolus</italic> Ausserer, 1871 and <italic>Harpactirella</italic> Purcell, 1902. Guadanucci &#x0026; Wendt (<xref ref-type="bibr" rid="cit0008">2014</xref>) and Zonstein (<xref ref-type="bibr" rid="cit0017">2018</xref>) conducted a taxonomic revision of Moroccan <italic>Ischnocolus</italic> species based on morphological data, and recorded the presence of three taxa: <italic>I. valentinus</italic> (Dufour, 1820), <italic>I. elongatus</italic> (Simon, 1873) and <italic>I. hancocki</italic> Smith, 1990. Whereas a considerable contribution to the taxonomy and biology of this genus in the Middle East has been recently published (see Montemor <italic>et al</italic>., <xref ref-type="bibr" rid="cit0013">2019</xref>), basic aspects of the biology, natural history and geographic distribution of the Moroccan species of <italic>Ischnocolus</italic> remain almost unknown. Similarly, no studies concerning the only species of the genus <italic>Harpactirella</italic> from Morocco, <italic>Harpactirella insidiosa</italic> (Denis, <xref ref-type="bibr" rid="cit0004">1960</xref>), have been published since its original description (Denis, <xref ref-type="bibr" rid="cit0004">1960</xref>) and subsequent taxonomic reassessment (Benoit, <xref ref-type="bibr" rid="cit0002">1965</xref>), being one of the least known species of Theraphosidae in the world.</p>
<p>The enigmatic spider and Moroccan endemic <italic>H. insidiosa</italic> was originally described in the monotypic genus <italic>Luphocemus</italic> Denis, <xref ref-type="bibr" rid="cit0004">1960</xref> [type species = <italic>Luphocemus insidiosus</italic>]<italic>.</italic> Denis (<xref ref-type="bibr" rid="cit0004">1960</xref>) described <italic>Luphocemus</italic> as follows: &#x201C;Pi&#x00E8;ces buccales et chelic&#x00E8;res des <italic>Leptopelma</italic> (= <italic>Ischnocolus sensu</italic> Denis, <xref ref-type="bibr" rid="cit0004">1960</xref>); Hauteur de bandeau inf&#x00E9;rieure au double du grand axe des yeux lat&#x00E9;raux ant&#x00E9;rieurs; Yeux post&#x00E9;rieurs sub&#x00E9;gaux; Intervalle des yeux lat&#x00E9;raux de chaque c&#x00F4;t&#x00E9; inf&#x00E9;rieur au diam&#x00E8;tre des yeux lat&#x00E9;raux post&#x00E9;rieurs; Article terminal des fili&#x00E8;res sup&#x00E9;rieures plus long que l&#x2019;article m&#x00E9;dian&#x201D; [Mouthparts and chelicerae as in <italic>Leptopelma</italic> (= <italic>Ischnocolus sensu</italic> Denis, <xref ref-type="bibr" rid="cit0004">1960</xref>); length of clypeus less than twice the distance between anterior-lateral eyes; sub-equal posterior eyes; distance between the lateral eyes of each side (anterior and posterior ones) less than the diameter of the posterior lateral eyes; distal segment of spinnerets larger than the medial segment]. Five years later, Benoit (<xref ref-type="bibr" rid="cit0002">1965</xref>) synonymized <italic>Luphocemus</italic> with <italic>Harpactirella</italic>, where it is currently allocated, based on some shared morphological traits between both taxa, namely a distal segment of spinnerets larger than the medial segment, and a similar morphology and relative position of the ocular tubercle and eyes.</p>
<p><italic>Harpactirella insidiosa</italic> is only known from two female specimens collected at two localities from Northern Morocco in the surroundings of Ben Slimane: one at Boulhaut, collected on 30 December 1951, and the second one at Sokhrat Nemra, collected on 18 March 1955 (Denis, <xref ref-type="bibr" rid="cit0004">1960</xref>). Males of <italic>H. insidiosa</italic> remain unknown.</p>
<p>In this article, we provide a new record based on the observation of two adult females of <italic>H. insidiosa</italic> far from their type locality. Both specimens were found in the Doukkala plain, 14 km southwest of Oualidia [approximate coordinates: 32&#x00B0;36&#x2019;N, 09&#x00B0;0&#x2019;W] (<xref ref-type="fig" rid="f0001">Fig. 1</xref>). The first one was observed under a rock on 29 December 2018 (<xref ref-type="fig" rid="f0002">Fig. 2</xref>), whereas the second recorded specimen was found within its burrow on 25 February 2020 (<xref ref-type="fig" rid="f0003">Fig. 3A</xref>). These findings extend the distribution of the species approximately 210 km southwest from the type locality. Measurements were taken from the first collected specimen, which had a total body length (carapace + abdomen) of approximately 20 mm, of which 9 mm correspond to the carapace (<xref ref-type="fig" rid="f0001">Fig. 1</xref>). Both specimens were compared with four topotypes collected from their burrows in Ben Slimane on 26 February 2020 (<xref ref-type="fig" rid="f0004">Fig. 4A&#x2013;B</xref>). All examined individuals present the traits described for the types of the species (Denis, <xref ref-type="bibr" rid="cit0004">1960</xref>). These topotypes exhibit similar colour patterns, except different brownish hues (<xref ref-type="fig" rid="f0004">Fig. 4C</xref>). The specimens from southern Oualidia slightly differ from the topotypes in the shape of the anterior-lateral eyes, which are more oval, and in their coloration, by showing a dorsal pattern of the opisthosoma with four distinctive oval dark marks, decreasing in size towards the posterior region, which are longitudinally arranged and bordered by additional transversal dark stripes. These marked transversal dark stripes on the abdominal sides are lacking in the topotype specimens which, in contrast, show a dotted abdominal pattern [Denis&#x2019; (<xref ref-type="bibr" rid="cit0004">1960</xref>) description only noted a brownish abdomen]. The ventral side of the abdomen is uniformly brown in the specimens from Oualidia (<xref ref-type="fig" rid="f0002">Fig. 2C</xref>). Burrows of both the Oualidia and topotype specimens were carefully excavated after checking for their entrances (<xref ref-type="fig" rid="f0003">Figs. 3A</xref>, <xref ref-type="fig" rid="f0004">4A</xref>). The structure of the burrows consisted of a vertical or sub-vertical tunnel 15&#x2013;25 cm deep, covered by silk, with a single small chamber at its end (pers.obs).</p>
<fig id="f0001">
<label>Fig. 1</label>
<caption><p>Map of Morocco showing known localities for <italic>Harpactirella insidiosa</italic>. Red dots represent literature records (Denis, <xref ref-type="bibr" rid="cit0004">1960</xref>). The yellow square corresponds to the new record at Doukkala plain.</p>
<p>Mapa de Marruecos con las localidades conocidas de <italic>Harpactirella insidiosa</italic>. Los puntos rojos representan registros bibliogr&#x00E1;ficos (Denis, <xref ref-type="bibr" rid="cit0004">1960</xref>). El cuadrado amarillo corresponde al nuevo registro de la llanura de Doukkala.</p></caption>
<graphic xlink:href="GRA2020_e113-g001.tif" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</fig>
<fig id="f0002">
<label>Fig. 2</label>
<caption><p>Adult female of <italic>Harpactirella insidiosa</italic> from Doukkala plain, Morocco. A: dorsal view; B: dorsolateral view; C: ventral view (Photographs AS-V).</p>
<p>Hembra adulta de <italic>Harpactirella insidiosa</italic> de la llanura de Doukkala, Marruecos. A: vista dorsal; B: vista dorsolateral; C: vista ventral (Fotograf&#x00ED;as AS-V).</p>
</caption>
<graphic xlink:href="GRA2020_e113-g002.tif" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</fig>
<fig id="f0003">
<label>Fig. 3</label>
<caption><p>Burrow entrance (A) and habitat of <italic>Harpactirella insidiosa</italic> (B) in Doukkala plain, Morocco (Photographs AS-V).</p>
<p>Entrada de la madriguera (A) y h&#x00E1;bitat de <italic>Harpactirella insidiosa</italic> (B) en Doukkala plain, Morocco (Fotograf&#x00ED;as AS-V).</p></caption>
<graphic xlink:href="GRA2020_e113-g003.tif" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</fig>
<fig id="f0004">
<label>Fig. 4</label>
<caption><p>Burrow entrance of a topotype specimen (A). Surroundings of Ben Slimane, type locality of <italic>Harpactirella insidiosa</italic> (B). Two topotypes of <italic>Harpactirella insidiosa</italic> (C) (Fotograf&#x00ED;as AS-V).</p>
<p>Entrada de la madriguera de un ejemplar topotipo (A). Alrededores de Ben Slimane, localidad tipo de <italic>Harpactirella insidiosa</italic> (B). Dos topotipos de <italic>Harpactirella insidiosa</italic> (A) (Fotograf&#x00ED;as AS-V).</p></caption>
<graphic xlink:href="GRA2020_e113-g004.tif" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</fig>
<p>The Doukkala plain has a semiarid climate (Escoriza &#x0026; Ben-Hassine, <xref ref-type="bibr" rid="cit0005">2019</xref>) and is mostly composed by limestone (Fakir <italic>et al</italic>., <xref ref-type="bibr" rid="cit0006">2002</xref>) with grasslands dominated by <italic>Asphodelus</italic> L. and scrubs of <italic>Chamaerops humilis</italic> L. and <italic>Olea europaea</italic> L., among others (<xref ref-type="fig" rid="f0003">Fig. 3B</xref>). The natural areas of the region are negatively impacted by agricultural development, <italic>Eucalyptus</italic> plantations, and urbanization in littoral areas (Hinckley <italic>et al</italic>., <xref ref-type="bibr" rid="cit0012">2016</xref>). Unfortunately, the lack of any biological information, including the ecological requirements of <italic>H. insidiosa</italic>, difficult the identification of potential threats that the species may be facing, preventing the proposal of specific conservation actions.</p>
<p>The finding of the presence of <italic>H. insidiosa</italic> in a semiarid climate, distinct from the warm Mediterranean climate predominant on the previously known localities, suggests that this species may be more broadly distributed that previously considered. However, molecular studies should be undertaken to confirm that the newly found population is conspecific with the northern populations of <italic>H. insidiosa</italic>. It should be borne in mind, however, that molecular studies in other mygalomorph taxa have revealed deep genetic divergences within single morphologically delimited species, usually corresponding to geographically restricted populations (Arnedo &#x0026; Ferr&#x00E1;ndez, <xref ref-type="bibr" rid="cit0001">2007</xref>; Hamilton <italic>et al</italic>., <xref ref-type="bibr" rid="cit0009">2011</xref>; Harvey <italic>et al</italic>., <xref ref-type="bibr" rid="cit0010">2015</xref>; Ferretti <italic>et al</italic>., <xref ref-type="bibr" rid="cit0006">2019</xref>). These studies suggest that the current diversity of mygalomorph spiders may have been grossly underestimated. Further integrative studies combining molecular and morphological, but also ecological and behavioural data may shed new light into the taxonomic status of the Doukkala population of <italic>H. insidiosa</italic>.</p>
<p>Molecular data may also contribute to resolve another important question posed by Moroccan <italic>Harpactirella</italic>. Except for the Moroccan <italic>H. insidiosa</italic>, the genus <italic>Harpactirella</italic> is exclusively composed of Afrotropical species, all of them endemic from South Africa, namely <italic>H. treleaveni</italic> Purcell, 1902, <italic>H. karroika</italic> Purcell, 1902, <italic>H. lightfooti</italic> Purcell, 1902, <italic>H. longipes</italic> Purcell, 1902, <italic>H. magna</italic> Purcell, 1903, <italic>H. helenae</italic> Purcell, 1903, <italic>H. domicola</italic> Purcell, 1903, <italic>H. schwarzi</italic> Purcell, 1904, <italic>H. lapidaria</italic> Purcell, 1908, <italic>H. spinosa</italic> Purcell, 1908, and <italic>H. overdijki</italic> Gallon, 2010 (World Spider Catalog, <xref ref-type="bibr" rid="cit0016">2020</xref>). If Benoit&#x2019;s (<xref ref-type="bibr" rid="cit0002">1965</xref>) inclusion of the Moroccan species in <italic>Harpactirella</italic> holds true, the resulting disjunct distribution showed by this genus represents an interesting biogeographic pattern that merits further investigation. Nevertheless, because of the rampant homoplasy of the few variable traits in an otherwise highly conserved phenotype (Raven, <xref ref-type="bibr" rid="cit0015">1985</xref>; Bond &#x0026; Opel, <xref ref-type="bibr" rid="cit0003">2002</xref>; Hamilton <italic>et al</italic>., <xref ref-type="bibr" rid="cit0009">2011</xref>), mygalomorph taxonomy has proven to be a challenging task (Hendrixson <italic>et al</italic>., <xref ref-type="bibr" rid="cit0011">2013</xref>; Harvey <italic>et al</italic>., <xref ref-type="bibr" rid="cit0010">2015</xref>; Ferretti <italic>et al</italic>., <xref ref-type="bibr" rid="cit0007">2019</xref>). The scrutiny of traditional taxonomic groups with new molecular information has revolutionized our understanding of the classification of mygalomorph spiders (e.g. Opatova <italic>et al</italic>., <xref ref-type="bibr" rid="cit0014">2019</xref>). In this regard, the synonymy of <italic>Luphocemus</italic> and <italic>Harpactirella</italic>, which was mostly based on two main morphological traits, may not resist a more detailed, molecular based re-examination.</p>
<p>The aforementioned observations highlight (1) the necessity of improving the taxonomic information on <italic>H. insidiosa</italic>, both at intra- and supra-specific levels, (2) the importance of compiling notes on its natural history, through carefully planned field work across the known localities and ecologically similar regions, (3) and the urgent need for measures to ensure conservation of this rare and poorly known species, inhabiting a vulnerable and highly disturbed region, outside natural protected areas.</p></body>
<back>
<ack>
<title>Acknowledgements</title>
<p>We are deeply grateful to Miguel Arnedo whose suggestions greatly improved the manuscript and to Miguel Angel Ferr&#x00E1;ndez for useful comments. We thank Javier Aznar Gonz&#x00E1;lez de Rueda for his help during fieldwork campaigns in Morocco between 2012 and 2019. Fieldwork was carried out with permits from the Haut Commissaire aux Eaux et For&#x00EA;ts et &#x00E0; la Lutte Contre la D&#x00E9;sertification (HCEFLCD /DLCDPN /DPRN/CFF N&#x00BA;19/2018).</p>
</ack>
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