ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2016 Magnolia Press

Zootaxa 4196 (2): 210–220 http://www.mapress.com/j/zt/

Article

http://doi.org/10.11646/zootaxa.4196.2.2

http://zoobank.org/urn:lsid:zoobank.org:pub:75113EF0-968D-45E3-9787-9D4FCF029737

Myrmeleon almohadarum sp. nov., from Spain and North Africa, with description

of the larva (Neuroptera Myrmeleontidae)

DAVIDE BADANO1,2, FERNANDO ACEVEDO3, ROBERTO A. PANTALEONI4,5 & VÍCTOR J. MONSERRAT31Istituto di Biologia Agroambientale e Forestale, Consiglio Nazionale delle Ricerche (IBAF–CNR), Via Salaria, km 29,300, I-00015

Monterotondo Scalo RM, Italy. E-mail: davide.badano@gmail.com2Centro Nazionale per lo Studio e la Conservazione della Biodiversità Forestale “Bosco Fontana", Strada Mantova 29, I-46045

Marmirolo MN, Italy3Departamento de Zoología y Antropología Física, Facultad de Biología, Universidad Complutense de Madrid, C/Jose Antonio

Novais, 2, 28040 Madrid, Spain. Email: facevedoramos@gmail.com; artmad@bio.ucm.es4Sezione di Entomologia, Dipartimento di Agraria, Università degli Studi, via Enrico De Nicola, I–07100 Sassari SS, Italy. E-mail:

pantaleo@uniss.it5Istituto per lo Studio degli Ecosistemi, Consiglio Nazionale delle Ricerche (ISE–CNR), Traversa la Crucca 3, Regione Baldinca, I–

07100 Li Punti SS, Italy. E-mail: r.pantaleoni@ise.cnr.it

Abstract

A new antlion (Neuroptera Myrmeleontidae Myrmeleontini), Myrmeleon almohadarum sp. nov., is described from south-

ern Spain and Tunisia. The new taxon is closely related to the mostly sympatric M. inconspicuus Rambur and

M. mariaemathildae Pantaleoni, Cesaroni & Nicoli Aldini but differing in body pattern, wing venation and larval chaeto-

taxy. The validity of the new species is also supported by a phylogenetic analysis based on COI sequences. The larva of

this new species is described and compared with congeners. M. almohadarum appears to be associated with sandy envi-

ronments.

Key words: Neuropterida, Myrmeleontini, antlion, Mediterranean, Iberian Peninsula, Tunisia

Introduction

The genus Myrmeleon Linnaeus is the most species-rich genus of the neuropteran family Myrmeleontidae, with around 180 described species, and the only one with a sub-cosmopolitan distribution, being absent only from oceanic islands and cold climates (Stange 2004). The larvae of this genus and closely related ones, i.e. EuroleonEsben-Petersen, attracted much attention being the most commonly encountered antlions building pit-traps (e.g.Gepp 2010; Devetak et al. 2012; Devetak & Arnett 2015). However, the considerable interest concerning the ethology and ecology of this genus is not counterbalanced by an adequate consideration of its taxonomy and phylogenetic relationships. The Euro-Mediterranean species were traditionally considered well known (e.g.Aspöck et al. 1980, 2001; Gepp 2010; Krivokhatsky 2011). Nevertheless, dedicated surveys conducted on the Italian islands and North Africa recently led to the discovery of new species of Myrmeleon (Pantaleoni et al. 2010; Pantaleoni & Badano 2012), implying an unnoticed diversity of this genus in the Mediterranean. During ongoing revisionary studies of Myrmeleontidae of the Iberian Peninsula (see Monserrat & Acevedo 2013) and North Africa, an undescribed species of Myrmeleon, similar to M. inconspicuus Rambur, but distinctly differing in body pattern, wing venation and larval chaetotaxy was found in southern Spain and Tunisia. Moreover, in order to reconstruct the relationships among the members of the M. inconspicuus species complex and to test the species boundaries in this morphologically homogeneous group, we performed a molecular phylogenetic analysis based on cytochrome c oxidase subunit I (COI). These interesting findings confirm that the Mediterranean region represents an important centre of speciation for this genus.

210 Accepted by B. Price: 6 Oct. 2016; published: 22 Nov. 2016

Materials and methods

Classical morphology. Most of the examined adults were obtained from larvae collected in the field. The larvae were reared in an insectary with a mean temperature of 25°C and kept in cylindrical containers filled with sand. The main offered prey were live larvae of mealworm, Tenebrio molitor Linnaeus (Coleoptera Tenebrionidae), and harvester ants, Messor barbarus Linnaeus (Hymenoptera Formicidae). The specimens were studied and measured with an Olympus® SZX7 and a Leica® MZ 9.5 stereomicroscopes provided with an optical micrometer. Photographs of both adults and larvae were taken with a digital Canon® EOS 600D camera equipped with Canon®

lens MP-E 65 mm; the resulting images were later processed with the stacking program Zerene® Stacker. Adult specimens were measured following the protocol of Pantaleoni et al. (2010) and Pantaleoni & Badano (2012): body length was measured from the vertex of the head to the tip of the abdomen; wing length was measured longitudinally from the insertion to the apex, while width was taken as the maximum width perpendicular to the length measurement line. A sample of 15 adults was measured: 6 males and 9 females. Larvae were measured according to Cesaroni et al. (2010) and Badano & Pantaleoni (2014): body length was measured from the head (excluding jaws) to the tip of abdomen; length of head capsule was measured ventrally from the distal to the proximal margin, head width was taken just below the eye tubercles, at the point of maximum width, mandible length was measured from apex to base. A sample of 8 third instar larvae was measured. The number of interdental mandibular setae is reported following the formula of Badano & Pantaleoni (2012): (a)(b)(c)(d), where a = average number of setae in the gap between mandible base and basal tooth; b = average number of setae in the gap between basal and median teeth; c = average number of setae in the gap between median and apical teeth; d = average number of setae in the gap between apical teeth and apex of the mandible. Male genitalia were macerated in 10% KOH (potaxium hydroxide) for several hours and later rinsed in acetic acid and water. Genitalia were finally stained in a saturated solution of Chlorazol Black in 95° ethanol and preserved in glycerol. Terminology

mainly follows Pantaleoni et al. (2010) and Badano & Pantaleoni (2014).

Examined collections. The study is based on 52 adult specimens (25 males, 27 females), preserved in the

following collections: Víctor Monserrat collection, Departamento de Zoología y Antropología Física,

Facultad de Biología, Universidad Complutense de Madrid, Madrid, Spain (VM); Roberto A. Pantaleoni

private collection, Istituto per lo Studio degli Ecosistemi, Consiglio Nazionale delle Ricerche, Sassari, Italy

(RAP); Davide Badano private collection, Taggia, Italy (DB). At least some specimens included in the type

series were previously identified as M. inconspicuus by Monserrat (1978) and Monserrat & Acevedo (2013),

these are marked here with an asterisk.

Distribution map. The map was compiled with the program ArcMap® 10.0 of the software ArcGis® 10.0.

Presence points were taken in a grid of 1x1 km2, in WGS84 international system coordinates.DNA sampling. COI sequences of the species of Myrmeleon were obtained from the GenBank database,

selecting specimens previously identified by D.B. and R.A.P., with a single exception (M. immanis, see Zhang and Whan 2014). The above mentioned sequences were previously extracted, amplified and deposited in the database by A. Schiaffino. Alignements were obtained using MUSCLE v.3.8 (Edgar 2004), implemented in Mesquite v.3.10 (Maddison and Maddison 2016), checking them visually with the latter software.

Phylogenetic reconstruction. The phylogenetic tree was inferred through a Bayesian approach, using MrBayes v.3.2.6 (Ronquist et al. 2012) on the CIPRES science gateway portal (Miller et al. 2010). The nucleotide substitution model GTR+I was selected as the best evolutionary model using ModelGenerator v.0.85 (Keane et al. 2006). Two parallel runs and four Markov Chains Monte Carlo (MCMC) were run for 5,000,000 generations and sampled every 1000th generation, with the first 10% of trees discarded as burn-in. Chain convergence was checked examining standard deviation of spit frequencies and using Tracer v.1.6 (Rambaut & Drummond 2007). The Bayesian consensus tree was then used as input for the Bayesian Poisson tree process algorithm (bPTP; Zhang et al. 2013) to infer species limits. In this case, the outgroup was removed to avoid biases on the results. The Markov chain was run for 100,000 generations with default settings for thinning and burn-in, finally checking convergence with the likelihood trace plot.

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Systematics

Myrmeleon almohadarum sp. nov.

Diagnosis. Small-sized Myrmeleon with a dark brown, variegated habitus (Figs 1, 2); pronotum with a diagnostic pattern (Fig. 3B); wings hyaline with dark-and-pale dashed veins (Fig. 4A); male hindwing with pilula axillaris; abdomen with conspicuous paler markings, creating an annulated pattern. Larva with an ochre habitus (Fig. 7), characterized by a particularly dense arrangement of digging setae on sternite IX (Fig. 8).

FIGURE 1. Myrmeleon almohadarum sp. nov., alcohol preserved male holotype and data labels.

FIGURE 2. Myrmeleon almohadarum sp. nov., female: habitus, dorsal (above) and lateral (below) view [Tunisia, Gammarth].

Description of the adult. Size. Average body length 20.43 mm (min-max 16.03–26.82); forewing male length 21.56 mm (19.20–23.33), female length 23.05 mm (19.84–26.5), ratio width/length (both sexes) 0.23; hind wing male length 19.57 mm (16.35–21.42), female length 20.81 mm (17.77–25.55), ratio width/length (both sexes) 0.22. General colouring. Dark brown with large ochre markings on thorax and abdomen (Fig. 2). Head. Vertex and occiput dark brown, with a contrasting ochre pattern (Fig. 3B). Frons dark brown with paler lateral margins. Ocular rim pale. Clypeus pale with fainted median marking. Labrum pale. Maxillary palpus brown. Labial palpus pale, with the distal segment fusiform and dark brown, palpimacula elliptical. Scape brown, paler distally, pedicellum dark brown, flagellum brown. (Fig. 3A). Thorax. Pronotum brown, with anterior and lateral margins pale; dorsal side with a complex pattern composed by: a narrow median pale stripe, a pair of poorly defined pale spots in the apical half and a pair of usually well contrasted pale spots in the basal half (Fig. 3B). Mesonotum and metanotum brown, lateral and ventral sclerite margins ochre, particularly evident on mesoscutum, mesoscutellum, metascutum and metascutellum (Fig. 2). Legs. Coxae brown in all legs (Fig. 2). Pro- and mesothoracic legs with extensive

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brown markings on the femur and tibia. Metathoracic leg with brown markings on the femur and the inner face of the tibia brown (Fig. 2). Tibial spurs as long as the first tarsomere. Wings. Relatively broad with a rounded apex (Fig. 4A). Membrane hyaline. Pterostigma distinct, proximally brown and distally whitish. Venation predominantly dark brown with alternating pale dashes. Radius sector of forewing with on average 5–7 (less frequently 8) crossveins, thus the cells of Rs are few and elongated. Cubital fork of forewing slightly more basal than Media posterior fork. Hind wing with on average 5 presectoral crossveins. Male hind wing equipped with pilula axillaris (Fig. 4A). Abdomen. Shorter than wings (Fig. 2). Tergites brown with large dorso-proximal ochre markings. Sternites brown with ventro-proximal pale markings. Interpleural membrane brown with paler markings. Ectoproct ochre. Male terminalia as in Fig. 5A, B; male genitalia, complex of gonocoxites 9+11 sensu Aspöck & Aspöck 2008 (gonarcus-paramere complex) as in Fig. 6. Female terminalia as in Fig. 5C, D.

FIGURE 3. Head and pronotum of M. inconspicuus species-group. A–B: Myrmeleon almohadarum sp. nov., A, head, frontal

view, B, head and pronotum, dorsal view [Tunisia, Gammarth]. C: Myrmeleon inconspicuus Rambur, head and pronotum,

dorsal view [Italy, Sardinia]. D: Myrmeleon mariaemathildae Pantaleoni, Cesaroni & Nicoli Aldini, head and pronotum, dorsal

view [Italy, Sardinia].

FIGURE 4. Wings of M. inconspicuus species-group. A: Myrmeleon almohadarum sp. nov., male wings [Tunisia, Gammarth].

B: Myrmeleon inconspicuus Rambur, forewing [Italy, Po Valley]. C: Myrmeleon mariaemathildae Pantaleoni, Cesaroni &

Nicoli Aldini, forewing [Italy, Sardinia].

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FIGURE 5. Myrmeleon almohadarum sp. nov.: male terminalia, A, lateral view, B, ventral view; female terminalia, C, lateral

view, D, ventral view [Tunisia, Gammarth].

FIGURE 6. Myrmeleon almohadarum sp. nov.: male genitalia, complex of gonocoxites 9 + gonocoxites 11 sensu Aspöck and

Aspöck, 2008 (= gonarcus-paramere complex), lateral (left) and ventral (right) view [Tunisia, Gammarth].

Variability. M. almohadarum sp. nov. is a relatively variable species in term of body pattern, like other congeners. Body colour varies from brown to dark brown but it is never pale ochre or blackish, like M. mariaemathildae and M. inconspicuus respectively. Some specimens are characterized by a partial fusion or fading of the pronotal pattern (e.g. specimen depicted in Fig. 2). The markings on the clypeus are also relatively variable.

Description of the third instar larva. Size. Average body length 9.20 mm; head length 1.78 mm (min-max 1.67–1.87), head width 1.53 mm (1.47–1.57), mandible length 1.83 mm (1.75–1.87), ratio head width/length 0.86, ratio mandible length/head length 1.03. General colouring. Pale brown, with a dark brown pattern, ventral side of the body paler, with conspicuous dark markings (Fig. 7). Dorsal side of the head capsule with dark brown markings on the clypeo-labrum and a posterior V-shaped, fainted marking, lateral side with dark brown markings. Ventral

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side pale, with a pair of large dark brown spots (Fig. 7). Mandible pale brown. Some preserved Spanish larvae are characterized by the presence of a dark marking in the median section of the mandible. Legs pale brown, unspotted. Body chaetotaxy black. Head. Sub-rectangular, longer than wide. Ocular tubercle sessile. Mandible as long as the head capsule, provided with 3 equidistant teeth not abruptly differentiated in size. Interdental mandibular setae: (5)(2)(2)(1). External margin with a fringe of long setae, dorsal and ventral side almost hairless except a few short setae near the margin. Labial palpus 4-segmented. Abdomen. VIII abdominal sternite with small odontoid processes, posterior margin with stout setae. IX abdominal sternite with many ventral digging setae, unevenly arranged or irregularly aligned in rows, mostly sub-equal in size and interspersed with smaller ones (Fig. 8). Rastra sessile, each bearing 4 digging setae, of which the external pairs are longer (Fig. 8).

Examined specimens. HOLOTYPE: Spain: Cádiz: Bolonia/ 30STE59/ 10 m/ ex larva 13.VII.2015 / F. Acevedo leg., 1 ♂ in alcohol [VM] (Fig. 1). PARATYPES. Spain: Almería: Las Casillas de Atochares, Rambla del Artal/ 210 m/ ex l. 20.VIII.2012 / F. Acevedo leg., 1 ♀ dry pinned [DB]; San Roque/ 300 m/ 2.IX.1993/ J. Ramírez leg., 1 ♀ dry pinned [VM]; Tabernas, Rambla Roja/ 360 m/ ex l. 16.VII.2012 / F. Acevedo leg., 1 ♂ dry pinned [DB]; Baleares: Ibiza, Las Salinas/ 10 m/ ex l. 3.VIII.2011 / V. J. Monserrat leg., 1 ♀ dry pinned [VM]; Cádiz: Barbate, Cerro del Pinar/ 130 m/ 20.VIII.1976 / V. J. Monserrat leg., 1♀* dry pinned [VM]; Pinar de la Duquesa 26.VIII.77 E.L. / V. Monserrat 1♀* [RAP]; Bolonia/ 10 m/ ex l. 13.VIII.2012 / F. Acevedo leg., 1♂* dry pinned [VM]; Bolonia/ 10 m/ ex l. 25.VIII.2012 / F. Acevedo leg., 1 ♂ dry pinned [VM]; Bolonia/ 10 m/ ex l. 13.VII.2015 / F. Acevedo leg., 1♂ [VM]; Bolonia/ 10 m/ ex l. 13.VII.2015 / F. Acevedo leg., 1♂ [VM]; Bonanza/ 18 m/ ex l. 2.VII.2012 / F. Acevedo leg., 1 ♂* dry pinned [DB]; Bonanza/ 18 m/ ex l. 3.VII.2012/ F. Acevedo leg., 1 ♂* dry pinned [VM]; Bonanza/ 18 m/ ex l. 23.VII.2012 / F. Acevedo leg., 1 ♂* dry pinned [VM]; Bonanza/ 18 m/ ex l. 13.VIII.2012 / F. Acevedo leg., 1 ♀* dry pinned [DB]; Cádiz, Castillo de San Sebastián/ 10 m/ 10.VII.1976 / V. J. Monserrat leg., 1 ♂* dry pinned [VM]; Caños de Meca/ 10 m/ ex l. 23.VII.2012 / F. Acevedo leg., 1 ♂* dry pinned [VM]; Caños de Meca/ 10 m/ ex l. 4.VIII.2012 / F. Acevedo leg., 1 ♂* dry pinned [VM]; Chiclana/ 10 m/ex l. 13.VII.2012 / F. Acevedo leg., 1 ♂* dry pinned [DB]; Playa de Los Lances/ 10 m/ ex l. 13.VIII.2012 / V. J. Monserrat leg., 1 ♂* dry pinned [DB]; Playa Los Lances/ 10 m/ ex l. 27.VIII.2012 / V. J. Monserrat leg., 1 ♀* dry pinned [DB]; Puerto Santa María, La Puntilla/ 10 m/ 29.VI.1976 / V. J. Monserrat leg., 1 ♂* dry pinned [VM]; Venta del Retín/ 40 m/ 5.VIII.1976 / I. Reviejo leg., 1 ♀* dry pinned [DB]; Huelva: Punta Umbría/ 10 m/ ex l. 16.VII.2012 / F. Acevedo leg., 1 ♀* dry pinned [VM]; Punta Umbría/ 10 m/ ex l. 4.VIII.2012 / F. Acevedo leg., 1 ♀* dry pinned [VM]; Jaén: El Centenillo/ 600 m/ ex l. 6.VII.1986/ V. J. Monserrat leg., 1 ♂[VM]; El Centenillo/ 600 m/ ex l. 10.VII.1986/ V. J. Monserrat leg., 1 ♂ [VM]; El Centenillo/ 600 m/ ex l. 10.VII.1986/ V. J. Monserrat leg., 1 ♀ [VM]; El Centenillo/ 600 m/ ex l. 11.VII.1986/ V. J. Monserrat leg., 1 ♀ [VM]; El Centenillo/ 600 m/ ex l. VIII.1986/ V. J. Monserrat leg., 1 ♀ [VM]; Málaga: Estepona, Sierra Bermeja, Rio Padrón/ 140 m/ 9.VII.2013 V. J. Monserrat leg., 1 ♂ dry pinned [DB].

Tunisia: Tunis, Gammarth / VII.2010 / INRGREF leg. / 1♀, ex larva, in alcohol [DB]; Tunis, Gammarth / VII.2010 / INRGREF leg. / 2♂, 2♀, ex larvae, in alcohol [RAP]; Forêt de Dar Chichou 36°57,733N 10°59,445E / VIII.2010 / INRGREF leg. 1♂, 4♀ ex larvae, in alcohol [RAP]; Al Sawasi (=Souassi) 35°21,953N 10°36,689E / VIII.2010 / INRGREF leg. 1♀ ex larva, in alcohol [RAP]; Tunis, Gammarth 36°55’10”N 10°17’38”E / 9.IV.2014 / R. A. Pantaleoni leg., 1 ♀, ex larva, dry pinned [DB]; Tunis, Gammarth 36°55’10”N 10°17’38”E / 9.IV.2014 / R. A. Pantaleoni leg. 1♂, 2♀, ex larvae, in alcohol [DB]; Tunis, Gammarth 36°55’10”N 10°17’38”E / 9.IV.2014 / R. A. Pantaleoni leg., 3♂, 2 ♀, ex larvae, in alcohol [RAP].

Larvae. Spain: Cádiz: Bolonia/ 10 m/ 24.V.2012 / F. Acevedo leg., 3 third instar larvae [VM]; Bolonia/ 10 m/ 5.IV.2015 / F. Acevedo leg., 2 third instar larvae [VM]; Bonanza/ 18 m/ 21.VI.2012 / F. Acevedo leg., 2 third instar larvae [VM]; Huelva: Punta Umbría/ 10 m/ 24.V.2012 / V. J. Monserrat leg., 3 third instar larvae [VM]; Málaga: Las Cañillas / 160 m/ 25.V.2012 / F. Acevedo leg., 6 third instar larvae [VM]; Monda, Llanos de Purla/ 370 m/ 18.VIII.2013/ V. J. Monserrat leg., 2 third instar larvae [VM].

Tunisia: Tunis, Gammarth 36°55’10”N 10°17’38”E / 9.IV.2014 / R. A. Pantaleoni leg., 2 third instar larvae, in alcohol [DB]; Tunis, Gammarth 36°55’10”N 10°17’38”E / 9.IV.2014 / R. A. Pantaleoni leg.,over 20 larvae, in alcohol [RAP]

Derivatio nominis. The name of this new species is a noun in the genitive case, referring to the Almohad dynasty, which dominated North Africa and the south of the Iberian Peninsula between the 12th and 13th centuries.

DNA taxonomy and phylogeny. The six putative species of the genus Myrmeleon, included in the analysed data set, were all supported by Bayesian posterior probabilities of 1.0 (Fig. 10). The Bayesian Poisson tree process algorithm applied on the Bayesian consensus tree also supported the existence of six species, reinforcing the morphology-based assumptions. The status of M. almohadarum as a new species was confirmed by a posterior probability of 0.97, given the input phylogeny and the bPTP model. The Bayesian phylogenetic analysis reconstructed M. almohadarum as sister to M. inconspicuus with a posterior probability of 1.0 (Fig. 10).

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FIGURE 7. Myrmeleon almohadarum sp. nov.: third instar larva, dorsal (above), ventral (middle) and lateral (below) view

[Tunisia, Gammarth].

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FIGURE 8. Myrmeleon almohadarum sp. nov.: third instar larva, detail of IX sternite, ventral view [Tunisia, Gammarth].

Ecological notes and distribution. In Tunisia, the pit-building larvae of M. almohadarum were collected in old coastal sand dunes with tree vegetation, including pine plantations. In Spain, besides the above mentioned environments, this species was also found in river banks and dry ephemeral riverbeds, suggesting that this antlion shares the same habitat preferences with M. inconspicuus (Nicoli Aldini 2007; Gepp 2010; Pantaleoni et al. 2010; Badano & Pantaleoni 2014) but it is more thermophilous. M. almohadarum is presently known from relatively few localities in Spain, mostly in the southernmost part of the Iberian Peninsula (Andalusia: Huelva, Jaén, Cádiz, Málaga and Almería). An isolated record from the Balearic Islands (Ibiza) is also known (Fig. 9). At least some of the numerous records of M. inconspicuus from the Iberian Peninsula (see list in Monserrat & Acevedo 2013), especially from its southernmost part and/or the Balearic Islands, very likely belong to the new species. In North Africa, this antlion was collected in Tunisia, but it is probably widespread along the south-western Mediterranean coast, in areas influenced by the Mediterranean climate and avoiding desert environments.

FIGURE 9. Myrmeleon almohadarum sp. nov.: distribution map.

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FIGURE 10. Rooted consensus tree of the phylogenetic relationships among analysed species of Myrmeleon based on a

Bayesian analysis of COI sequences. Average branch lengths proportional to numbers of substitutions per site under the GTR+I

substitution model. Bayesian posterior probabilities are shown above each branch; support values for within-species

relationships are not shown for very short branches. Species nodes identified by bPTP are highlighted by red terminal branches.

Specimen codes are GenBank accessions.

Comparative notes. The Myrmeleon inconspicuus-group of species, firstly delimited by Pantaleoni et al. (2010), includes the Turano-European M. inconspicuus, the Mediterranean M. mariaemathildae and the Asiatic-E-European M. immanis Walker. These species differ from other Palaearctic congeners in a set of adult characters: forewing Cubital fork more basal than Media posterior fork (Fig. 4), male hind wing with pilula axillaris and male genitalia with a large, lamellar mediuncus (Fig. 6). Moreover, the larvae of this species-group are characterized by the IX abdominal sternite with the anterior row of digging setae composed by at least 6 bristles (Badano & Pantaleoni 2014). M. almohadarum also fits within this species group, based on adult and larval morphology and the DNA-based phylogenetic reconstruction. The new species and M. immanis are easily distinguished by the unmistakable habitus of the latter (i.e. blackish body colour and yellowish, hyaline wings with pointed apex). M.inconspicuus is also darker than M. almohadarum and it is characterized by a blackish pronotum with a thin median line and an isolated pale spot per side (Fig. 3C). On the other hand, M. mariaemathildae is a highly variable species, with distinct pale and dark morphs, though most specimens are pale ochre. The pronotum of M.mariaemathildae is usually characterized by large pale markings and paired dark areas, with a large median, pale stripe connected with a pair of posterior spots (Fig. 3D), though in very dark specimens the pronotum closely resembles that of M. inconspicuus, see Pantaleoni et al. (2010) for further details. M. almohadarum is best distinguished from these closely related species due to the overall brown habitus and the dark brown pronotum with two pale markings per side connected to a median stripe (but see Variability). The new species is usually characterized by the Radius sector of forewing with about 5–7 transversal crossveins, therefore the cells of the Rs are few and comparatively long (Fig. 4A). In other species (including the other members of the M. inconspicuusgroup) the Rs crossveins are usually 10 or more, thus the cells appear more numerous but shorter (Fig. 4B, C). M. almohadarum closely resembles the African species M. caliginosus Hölzel & Ohm. The most obvious difference, besides genitalia, between these species is represented by the presence of pilula axillaris on male hind wing, always lacking in M. caliginosus. Moreover, M. caliginosus is overall darker, with pronotal spots well separated and not connected by a median line.

The larva of M. almohadarum is very similar in body pattern and relative proportions to those of M. inconspicuus and M. mariaemathildae (see Badano & Pantaleoni 2014). The larva of the new species is characterized by the IX sternite with the anterior row of digging setae comprising numerous and irregularly

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arranged bristles, mostly of similar size (Fig. 8). M. inconspicuus has a lower number of setae on the anterior row, while M. mariaemathildae is recognizable due to its combination of large setae disorderly interspersed with smaller ones (Badano & Pantaleoni 2014).

Acknowledgements

This contribution is part of the coordinated project of I+D+i: "Fauna Ibérica (Neuroptera) (CGL2010-22267-C07-05)". Special thanks to Angela Schiaffino (ISE–CNR) for her invaluable help in DNA extraction and sequencing.

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AbstractIntroductionMaterials and methodsSystematicsMyrmeleon almohadarum sp. nov.AcknowledgementsReferences

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