4196 (2): 210 220 Article …...(RAP); Davide Badano private collection, Taggia, Italy (DB). At...

210 Accepted by B Price 6 Oct 2016 published 22 Nov 2016

ZOOTAXA

ISSN 1175-5326 (print edition)

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

Zootaxa 4196 (2) 210ndash220

httpwwwmapresscomjztArticle

httpdoiorg1011646zootaxa419622

httpzoobankorgurnlsidzoobankorgpub75113EF0-968D-45E3-9787-9D4FCF029737

Myrmeleon almohadarum sp nov from Spain and North Africa with description

of the larva (Neuroptera Myrmeleontidae)

DAVIDE BADANO12 FERNANDO ACEVEDO3 ROBERTO A PANTALEONI45 amp VIacuteCTOR J MONSERRAT3

1Istituto di Biologia Agroambientale e Forestale Consiglio Nazionale delle Ricerche (IBAFndashCNR) Via Salaria km 29300 I-00015

Monterotondo Scalo RM Italy E-mail davidebadanogmailcom2Centro Nazionale per lo Studio e la Conservazione della Biodiversitagrave Forestale ldquoBosco Fontana Strada Mantova 29 I-46045

Marmirolo MN Italy3Departamento de Zoologiacutea y Antropologiacutea Fiacutesica Facultad de Biologiacutea Universidad Complutense de Madrid CJose Antonio

Novais 2 28040 Madrid Spain Email facevedoramosgmailcom artmadbioucmes4Sezione di Entomologia Dipartimento di Agraria Universitagrave degli Studi via Enrico De Nicola Indash07100 Sassari SS Italy E-mail

pantaleounissit5Istituto per lo Studio degli Ecosistemi Consiglio Nazionale delle Ricerche (ISEndashCNR) Traversa la Crucca 3 Regione Baldinca Indash

07100 Li Punti SS Italy E-mail rpantaleoniisecnrit

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 amp 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 ie Euroleon

Esben-Petersen attracted much attention being the most commonly encountered antlions building pit-traps (eg

Gepp 2010 Devetak et al 2012 Devetak amp 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 (eg

Aspoumlck 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 amp Badano 2012) implying an unnoticed diversity of this genus in the Mediterranean During ongoing revisionary studies of Myrmeleontidae of the Iberian Peninsula (see Monserrat amp 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

Zootaxa 4196 (2) copy 2016 Magnolia Press middot 211MYRMELEON ALMOHADARUM SP NOV

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 25degC 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 Olympusreg SZX7 and a Leicareg MZ 95 stereomicroscopes provided with an optical micrometer Photographs of both adults and larvae were taken with a digital Canonreg EOS 600D camera equipped with Canonreg

lens MP-E 65 mm the resulting images were later processed with the stacking program Zerenereg Stacker Adult specimens were measured following the protocol of Pantaleoni et al (2010) and Pantaleoni amp 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 amp 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 amp 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 95deg ethanol and preserved in glycerol Terminology

mainly follows Pantaleoni et al (2010) and Badano amp Pantaleoni (2014)

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

following collections Viacutector Monserrat collection Departamento de Zoologiacutea y Antropologiacutea Fiacutesica

Facultad de Biologiacutea 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 amp Acevedo (2013)

these are marked here with an asterisk

Distribution map The map was compiled with the program ArcMapreg 100 of the software ArcGisreg 100

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

selecting specimens previously identified by DB and RAP 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 v38 (Edgar 2004) implemented in Mesquite v310 (Maddison and Maddison 2016) checking them visually with the latter software

Phylogenetic reconstruction The phylogenetic tree was inferred through a Bayesian approach using MrBayes v326 (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 v085 (Keane et al 2006) Two parallel runs and four Markov Chains Monte Carlo (MCMC) were run for 5000000 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 v16 (Rambaut amp 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 100000 generations with default settings for thinning and burn-in finally checking convergence with the likelihood trace plot

BADANO ET AL212 middot Zootaxa 4196 (2) copy 2016 Magnolia Press

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 2043 mm (min-max 1603ndash2682) forewing male length 2156 mm (1920ndash2333) female length 2305 mm (1984ndash265) ratio widthlength (both sexes) 023 hind wing male length 1957 mm (1635ndash2142) female length 2081 mm (1777ndash2555) ratio widthlength (both sexes) 022 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


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 5ndash7 (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 Aspoumlck amp Aspoumlck 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 AndashB 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 amp 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 amp

Nicoli Aldini forewing [Italy Sardinia]


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 Aspoumlck and

Aspoumlck 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 (eg specimen depicted in Fig 2) The markings on the clypeus are also relatively variable

Description of the third instar larva Size Average body length 920 mm head length 178 mm (min-max 167ndash187) head width 153 mm (147ndash157) mandible length 183 mm (175ndash187) ratio head widthlength 086 ratio mandible lengthhead length 103 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


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 Caacutediz Bolonia 30STE59 10 m ex larva 13VII2015 F Acevedo leg 1 in alcohol [VM] (Fig 1) PARATYPES Spain Almeriacutea Las Casillas de Atochares Rambla del Artal 210 m ex l 20VIII2012 F Acevedo leg 1 dry pinned [DB] San Roque 300 m 2IX1993 J Ramiacuterez leg 1 dry pinned [VM] Tabernas Rambla Roja 360 m ex l 16VII2012 F Acevedo leg 1 dry pinned [DB] Baleares Ibiza Las Salinas 10 m ex l 3VIII2011 V J Monserrat leg 1 dry pinned [VM] Caacutediz Barbate Cerro del Pinar 130 m 20VIII1976 V J Monserrat leg 1 dry pinned [VM] Pinar de la Duquesa 26VIII77 EL V Monserrat 1 [RAP] Bolonia 10 m ex l 13VIII2012 F Acevedo leg 1 dry pinned [VM] Bolonia 10 m ex l 25VIII2012 F Acevedo leg 1 dry pinned [VM] Bolonia 10 m ex l 13VII2015 F Acevedo leg 1 [VM] Bolonia 10 m ex l 13VII2015 F Acevedo leg 1 [VM] Bonanza 18 m ex l 2VII2012 F Acevedo leg 1 dry pinned [DB] Bonanza 18 m ex l 3VII2012 F Acevedo leg 1 dry pinned [VM] Bonanza 18 m ex l 23VII2012 F Acevedo leg 1 dry pinned [VM] Bonanza 18 m ex l 13VIII2012 F Acevedo leg 1 dry pinned [DB] Caacutediz Castillo de San Sebastiaacuten 10 m 10VII1976 V J Monserrat leg 1 dry pinned [VM] Cantildeos de Meca 10 m ex l 23VII2012 F Acevedo leg 1 dry pinned [VM] Cantildeos de Meca 10 m ex l 4VIII2012 F Acevedo leg 1 dry pinned [VM] Chiclana 10 mex l 13VII2012 F Acevedo leg 1 dry pinned [DB] Playa de Los Lances 10 m ex l 13VIII2012 V J Monserrat leg 1 dry pinned [DB] Playa Los Lances 10 m ex l 27VIII2012 V JMonserrat leg 1 dry pinned [DB] Puerto Santa Mariacutea La Puntilla 10 m 29VI1976 V J Monserrat leg 1 dry pinned [VM] Venta del Retiacuten 40 m 5VIII1976 I Reviejo leg 1 dry pinned [DB] Huelva PuntaUmbriacutea 10 m ex l 16VII2012 F Acevedo leg 1 dry pinned [VM] Punta Umbriacutea 10 m ex l 4VIII2012 F Acevedo leg 1 dry pinned [VM] Jaeacuten El Centenillo 600 m ex l 6VII1986 V J Monserrat leg 1 [VM] El Centenillo 600 m ex l 10VII1986 V J Monserrat leg 1 [VM] El Centenillo 600 m ex l10VII1986 V J Monserrat leg 1 [VM] El Centenillo 600 m ex l 11VII1986 V J Monserrat leg 1

[VM] El Centenillo 600 m ex l VIII1986 V J Monserrat leg 1 [VM] Maacutelaga Estepona Sierra BermejaRio Padroacuten 140 m 9VII2013 V J Monserrat leg 1 dry pinned [DB]

Tunisia Tunis Gammarth VII2010 INRGREF leg 1 ex larva in alcohol [DB] Tunis Gammarth VII2010 INRGREF leg 2 2 ex larvae in alcohol [RAP] Forecirct de Dar Chichou 36deg57733N 10deg59445E VIII2010 INRGREF leg 1 4 ex larvae in alcohol [RAP] Al Sawasi (=Souassi) 35deg21953N 10deg36689E VIII2010 INRGREF leg 1 ex larva in alcohol [RAP] Tunis Gammarth 36deg55rsquo10rdquoN 10deg17rsquo38rdquoE 9IV2014 R A Pantaleoni leg 1 ex larva dry pinned [DB] Tunis Gammarth 36deg55rsquo10rdquoN 10deg17rsquo38rdquoE 9IV2014 RA Pantaleoni leg 1 2 ex larvae in alcohol [DB] Tunis Gammarth 36deg55rsquo10rdquoN 10deg17rsquo38rdquoE 9IV2014 RA Pantaleoni leg 3 2 ex larvae in alcohol [RAP]

Larvae Spain Caacutediz Bolonia 10 m 24V2012 F Acevedo leg 3 third instar larvae [VM] Bolonia 10 m5IV2015 F Acevedo leg 2 third instar larvae [VM] Bonanza 18 m 21VI2012 F Acevedo leg 2 third instarlarvae [VM] Huelva Punta Umbriacutea 10 m 24V2012 V J Monserrat leg 3 third instar larvae [VM] MaacutelagaLas Cantildeillas 160 m 25V2012 F Acevedo leg 6 third instar larvae [VM] Monda Llanos de Purla 370 m18VIII2013 V J Monserrat leg 2 third instar larvae [VM]

Tunisia Tunis Gammarth 36deg55rsquo10rdquoN 10deg17rsquo38rdquoE 9IV2014 R A Pantaleoni leg 2 third instar larvae inalcohol [DB] Tunis Gammarth 36deg55rsquo10rdquoN 10deg17rsquo38rdquoE 9IV2014 R A Pantaleoni legover 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 10 (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 097 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 10 (Fig 10)


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

[Tunisia Gammarth]


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 amp 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 Jaeacuten Caacutediz Maacutelaga and Almeriacutea) 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 amp Acevedo 2013) especially from its southernmost part andor 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


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 amp 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 (ie 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 5ndash7 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 inconspicuus

group) 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 Houmllzel amp 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 amp 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


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 amp Pantaleoni 2014)

Acknowledgements

This contribution is part of the coordinated project of I+D+i Fauna Ibeacuterica (Neuroptera) (CGL2010-22267-C07-05) Special thanks to Angela Schiaffino (ISEndashCNR) for her invaluable help in DNA extraction and sequencing

References

Aspoumlck U amp Aspoumlck H (2008) Phylogenetic relevance of the genital sclerites of Neuropterida (Insecta Holometabola)

Systematic Entomology 33 97ndash127

httpdxdoiorg101111j1365-3113200700396x

Aspoumlck H Aspoumlck U amp Houmllzel H (1980) Die Neuropteren Europas Goecke and Evers Krefeld Germany 495 + 355 pp

Aspoumlck H Houmllzel H amp Aspoumlck U (2001) Kommentierter Katalog der Neuropterida (Insecta Raphidioptera Megaloptera

Neuroptera) der Westpalaumlarktis Denisia 2 1ndash606

Badano D amp Pantaleoni R A (2014) The larvae of European Myrmeleontidae (Neuroptera) Zootaxa 3762 1ndash71

httpdxdoiorg1011646zootaxa376211

Cesaroni C Nicoli Aldini R amp Pantaleoni RA (2010) The larvae of Gymnocnemia variegata (Schneider 1845) and

Megistopus flavicornis (Rossi 1790) (Neuroptera Myrmeleontidae) a comparative description In Devetak D

Lipovšek S amp Arnett AE (Eds) Proceedings of the 10th International Symposium on Neuropterology (22ndash25 June

2008 Piran Slovenia) University of Maribor Maribor Slovenia pp 135ndash144

Devetak D amp Arnett AE (2015) Preference of antlion and wormlion larvae (Neuroptera Myrmeleontidae Diptera

Vermileonidae) for substrates according to substrate particle size European Journal of Entomology 112 500ndash509

httpdxdoiorg1014411eje2015052

Devetak D Novak T amp Janžekovič F (2012) Effect of substrate density on behaviour of antlion larvae (Neuroptera

Myrmeleontidae) Acta Oecologica 43 1ndash7

httpdxdoi1011861471-2148-6-29

Edgar RC (2004) MUSCLE multiple sequence alignment with high accuracy and high throughput Nucleic Acids Research

32 1792ndash1797

Gepp J (2010) Ameisenloumlwen und Ameisenjungfern Myrmeleontidae Eine weltweite Betrachtung unter besonderer

Beruumlcksichtigung Mitteleuropas 3 neubearbeitete Auflage Die Neue Brehm-Buumlcherei 589 Westarp Wissenschaften-

Verlagsgesellschaft Hohenwarsleben Germany 168 pp

Keane TM Creevey CJ Pentony MM Naughton TJ amp Mclerney JO (2006) Assessment of methods for amino acid

matrix selection and their use on empirical data shows that ad hoc assumptions for choice of matrix are not justified BMC

Evolutionary Biology 6 29

httpdxdoi1011861471-2148-6-29

Krivokhatsky VA (2011) Antlions (Neuroptera Myrmeleontidae) of Russia KMK Saint Petersburg Russia 334 pp [in

Russian]

Maddison WP amp Maddison DR (2016) Mesquite A Modular System for Evolutionary Analysis Version 310 URL http

mesquiteprojectorg [accessed on 5 May 2016]

Miller MA Pfeiffer WT amp Schwartz T (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic

trees Proceedings of the Gateway Computing Environments Workshop (GCE) 1ndash8

Monserrat VJ (1978) Primera contribucioacuten al conocimiento de los Neuroacutepteros de Caacutediz (Insecta Neuroptera Planipennia)

Boletiacuten Real Sociedad Espantildeola Historia Natural (Biol) 76 57ndash70

Monserrat VJ amp Acevedo F (2013) Los mirmeleoacutenidos (hormigas-leoacuten) de la Peniacutensula Ibeacuterica e Islas Baleares (Insecta

Neuropterida Neuroptera Myrmeleontidae) Graellsia 69 283ndash321

httpdxdoiorg103989graellsia2013v69098

Nicoli Aldini R (2007) Observations on the larval morphology of the antlion Myrmeleon bore (Tjeder 1941) (Neuroptera

Myrmeleontidae) and its life cycle in the Po Valley (northern Italy) In Pantaleoni RA Letardi A amp Corazza C (Eds)

Proceedings of the Ninth International Symposium on Neuropterology (20ndash23 June 2005 Ferrara Italy) Annali del

Museo Civico di Storia Naturale di Ferrara 8 59ndash66

Pantaleoni RA amp Badano D (2012) Myrmeleon punicanus n sp a new pit-building antlion from Sicily and Pantelleria

Bulletin of Insectology 65 139ndash148

Pantaleoni RA Cesaroni C amp Nicoli Aldini R (2010) Myrmeleon mariaemathildae n sp a new Mediterranean pit-

building antlion (Neuropterida Myrmeleontidae) Bulletin of Insectology 63 91ndash98


Rambaut A amp Drummond AJ (2007) Tracer v16 URL httpbeastbioedacukTracer [accessed on 1 September 2016]

Ronquist F Teslenko M van der Mark P Ayres D Darling A Houmlhna S Larget B Liu L Suchard MA amp

Huelsenbeck JP (2012) MrBayes 32 efficient Bayesian phylogenetic inference and model choice across a large model

space Systematic Biology 61 539ndash542

Stange LA (2004) A systematic catalog bibliography and classification of the world antlions (Insecta Neuroptera

Myrmeleontidae) Memoirs of the American Entomological Institute 74 1ndash565

Zhang J Kapli P Pavlidis P amp Stamatakis S (2013) A general species delimitation method with applications to

phylogenetic placements Bioinformatics 29 2869ndash2876

Zhang J amp Wang X-L (2014) The complete mitochondrial genome of Myrmeleon immanis Walker 1853 (Neuroptera

Myrmeleontidae) Mitochondrial DNA 27 1439ndash1440


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 25degC 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 Olympusreg SZX7 and a Leicareg MZ 95 stereomicroscopes provided with an optical micrometer Photographs of both adults and larvae were taken with a digital Canonreg EOS 600D camera equipped with Canonreg

lens MP-E 65 mm the resulting images were later processed with the stacking program Zerenereg Stacker Adult specimens were measured following the protocol of Pantaleoni et al (2010) and Pantaleoni amp 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 amp 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 amp 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 95deg ethanol and preserved in glycerol Terminology

mainly follows Pantaleoni et al (2010) and Badano amp Pantaleoni (2014)

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

following collections Viacutector Monserrat collection Departamento de Zoologiacutea y Antropologiacutea Fiacutesica

Facultad de Biologiacutea 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 amp Acevedo (2013)

these are marked here with an asterisk

Distribution map The map was compiled with the program ArcMapreg 100 of the software ArcGisreg 100

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

selecting specimens previously identified by DB and RAP 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 v38 (Edgar 2004) implemented in Mesquite v310 (Maddison and Maddison 2016) checking them visually with the latter software

Phylogenetic reconstruction The phylogenetic tree was inferred through a Bayesian approach using MrBayes v326 (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 v085 (Keane et al 2006) Two parallel runs and four Markov Chains Monte Carlo (MCMC) were run for 5000000 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 v16 (Rambaut amp 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 100000 generations with default settings for thinning and burn-in finally checking convergence with the likelihood trace plot


Systematics



































[Tunisia Gammarth]




















Acknowledgements


References



httpdxdoiorg101111j1365-3113200700396x















httpdxdoi1011861471-2148-6-29


32 1792ndash1797







httpdxdoi1011861471-2148-6-29


Russian]






























Systematics



































[Tunisia Gammarth]




















Acknowledgements


References



httpdxdoiorg101111j1365-3113200700396x















httpdxdoi1011861471-2148-6-29


32 1792ndash1797







httpdxdoi1011861471-2148-6-29


Russian]


























































[Tunisia Gammarth]




















Acknowledgements


References



httpdxdoiorg101111j1365-3113200700396x















httpdxdoi1011861471-2148-6-29


32 1792ndash1797







httpdxdoi1011861471-2148-6-29


Russian]
















































Acknowledgements


References



httpdxdoiorg101111j1365-3113200700396x















httpdxdoi1011861471-2148-6-29


32 1792ndash1797







httpdxdoi1011861471-2148-6-29


Russian]





























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