Taxonomic notes on the antlion tribe Myrmeleontini Latreille (Neuroptera, Myrmeleontidae, Myrmeleontinae) from Pakistan, with description of a new species

. A new species of the myrmeleontine antlion genus Baliga Navás, 1912 (Neuroptera: Myrmeleontidae), Baliga kashmirensis sp. nov., from Azad Kashmir and Khyber Pakhtunkhwa Province of Pakistan is described and illustrated, representing the ﬁ rst record of Baliga from Pakistan. Three species of Myrmeleon Linnaeus, 1767 are re-described: M. hyalinus hyalinus Olivier, 1811, M. tenuipennis Rambur, 1842, and M. trivialis Gerstaecker, 1885. Myrmeleon bimaculatus Yang, 1999 syn. nov. originally described from China is considered to be a junior synonym of Myrmeleon tenuipennis Rambur, 1842. In addition, an annotated catalogue of all species of Myrmeleon known from Pakistan along with their distribution map, taxonomical notes and updated identi ﬁ cation key to known genera and species are provided. Taxonomic notes on the antlion tribe Myrmeleontini Latreille (Neuroptera, Myrmeleontidae, Myrmeleontinae) from Pakistan, with description of a new species. European Journal of

An interesting result in the recent phylogenetic analysis of Myrmeleontidae by Machado et al. (2019) suggested that Baliga, Hagenomyia, together with six genera (Australeon, Dictyoleon, Euroleon, Kirghizoleon, Megistoleon, and Weeleus) are potential synonyms of Myrmeleon. The results of Machado et al. (2019) could also be interpreted that Myrmeleon, as currently circumscribed, is massively paraphyletic. Despite the recent efforts on phylogeny and the larval taxonomy of antlions (Badano et al. 2017b;Machado et al. 2019), there are a few studies that inferred morphological affi nities which used adult characters to infer antlion phylogeny (Stange 1994;Badano et al. 2017cBadano et al. , 2018Machado & Oswald 2020). The larvae of all known genera of Myrmeleontini are pit-builders and exhibit a striking similar morphology. For example, the larvae of Myrmeleon and Euroleon are remarkably similar, but their adults can be distinguished based on wing venational characters (Mansell 1996;Stange 2004;Badano & Pantaleoni 2014;Badano et al. 2017b). In fact, many myrmeleontine genera, especially Baliga and Hagenomyia, are based on variable morphological characters, questioning the affi nities of several taxa. Navás (1912b) originally described Baliga based on the presence of interconnected crossveins in the costal area proximal to the forewing pterostigma. Later, Esben-Petersen (1913) synonymized this genus with Hagenomyia and remarked that the description of Baliga solely based on the presence of interconnected crossveins is not justifi ed. This view was later followed by Markl (1954), Kuwayama (1962), Oswald & Penny (1991), Ghosh (2000), Bao et al. (2007), Wang et al. (2018), and Yang et al. (2018). However, Stange & Wang (1998), Stange (2004), and Hayashi et al. (2020) considered Baliga as a valid genus based on the shorter length of anterior gonocoxites 8 than posterior gonocoxites 8 (= anterior and posterior gonapophyses in Stange 2004) in the female genitalia (these two sclerites are nearly equal in length in Hagenomyia).
In this paper, we fi rst report Baliga from Pakistan based on a new species, namely Baliga kashmirensis sp. nov. Then, we summarize the present information of Myrmeleontini from Pakistan and add some new fi ndings of this tribe from our recent fi eld surveys in the northern parts of Pakistan. Finally, we also use molecular data, i.e., the partial DNA sequences of the mitochondrial COI and 16S rRNA genes, to verify the validity of the new species herein described. A dichotomous key as well as a distribution map are provided for Baliga and Myrmeleon species in Pakistan.

Taxonomy
The adult specimens were collected at day time (around 10.00 am in the morning to evening at 6 pm) by using a sweeping net along the mountainous regions in Azad Kashmir, Khyber Pakhtunkhwa and Punjab provinces of Pakistan. Samples were preserved in 95% ethanol or pinned. The specimens examined are deposited in the Entomological Museum, China Agricultural University (CAU), Beijing, China and the following institutes in Pakistan: Pakistan Museum of Natural History (PMNH), and the National Insect Museum (NIM), Islamabad. A male of Baliga sagax (Walker, 1853) (collecting information: "CHINA: Fujian, Xiamen, Siming District, Huandao Road, Wanyuepo, 0 m, 25.vi.2021, leg. Yuchen Zheng") was used for morphological comparison with Baliga kashmirensis sp. nov.
Photographs of the adult habitus were taken with a Nikon D800 or D850 digital camera with a Nikon Micro-Nikkor 105 mm lens, while the genital photographs were taken with a Canon 7D Mark II or D850 digital camera with a Nikon SMZ18 microscope. Photographs were cleaned up and laid out with Helicon focus (ver. 6.7.1), and Adobe Photoshop CS 6.0. The distribution map was prepared with ArcGIS 10.5 software (Esri, Redlands, CA) by using the original base map of Pakistan.

Molecular identifi cation
Representatives of species of Baliga and Myrmeleon sampled from Pakistan and the published data of the Japanese and Chinese species of these two genera (Hayashi et al. 2020) were included for the molecular identifi cation of the species from Pakistan. Moreover, the available COI data of six species of Myrmeleon from Pakistan (Akhtar 2018), Myrmeleon hyalinus from Egypt and Greece, and two subspecies of Myrmeleon hyalinus Olivier, 1811 from Azerbaijan retrieved from the NCBI GenBank were also included for molecular analysis. Since no 16S rRNA gene sequences were available in GenBank for those six species of Myrmeleon from Pakistan and two subspecies of Myrmeleon hyalinus from Azerbaijan, we used solely the 16S RNA gene sequences to generate the phylogenetic tree for species of Baliga and Myrmeleon sampled in the present study and those from the Chinese and Japanese species. Norfolius howensis (Tillyard, 1917) and Nymphes myrmeleonoides Leach, 1814 (Neuroptera: Nymphidae) were selected as outgroups (Winterton et al. 2010). The species list, along with their corresponding GenBank accession numbers and collecting data, are provided in Table 1.
The adult specimens were stored in 95% ethanol and refrigerated at -20°C. The genomic DNA was extracted from the thoracic muscle or several legs of each individual sample preserved in ethanol. The samples were incubated in the extraction buffer/proteinase-K mixture at 56°C for 12 h. We used the DNeasy Blood & Tissue kit (QIAGEN, Beijing, China) or TIANamp Micro DNA Kit (TIANGEN BIOTECH CO., LTD, Beijing, China), following manufacturers' instructions. Partial sequences of two mitochondrial genes, i.e., COI and 16S rRNA, were amplifi ed and sequenced. PCR was carried out in a Bio-Rad/T100™ Thermal Cycler (Hercules, CA, USA) by using an AccuPower PCR Premix (Bioneer, Daejeon, Korea) with 12.5 L in a reaction volume of 25 L, which included 1.0 L DNA templet, 1.0 L forward primer, 1.0 L reverse primer and 9.5 L distilled water.

Taxonomy
Tribe Myrmeleontini Latreille, 1802 Key to genera of Myrmeleontini from Pakistan 1. Antenna slightly swollen; wings generally broad; forewing costal area usually with interconnected crossveins proximal to pterostigmal area ( Fig. 2A-B

Diagnosis
Baliga is quite similar to Hagenomyia and Myrmeleon but can be distinguished from Hagenomyia by the female anterior gonocoxites 8 relatively shorter than posterior gonocoxites 8 (these two female genital sclerites are nearly equal length in Hagenomyia) and from Myrmeleon by the presence of interconnected crossveins in the costal area of forewing (these interconnected crossveins are absent in Myrmeleon).

Distribution
Baliga currently includes 17 described species, predominantly distributed in the Oriental (12 species) and Palaearctic regions (4 spp.: China, Japan, and Korea) with a single species in Australia (Queensland). It is widely distributed in the Oriental region: Bangladesh, China, India, Myanmar, Sri Lanka, Vietnam, and the main islands of Indonesia, Malaysia, Micronesia, Philippines, Japan, and Korea (Ghosh 2000;Stange 2004;Bao et al. 2007;Hayashi et al. 2020

Diagnosis
Larger-sized species (forewing length: 34.5-44.8 mm), superfi cially resembling to Baliga sagax (Walker, 1853) based on similar yellow markings on vertex but can be distinguished by the yellow pronotum, with a pair of well separated median longitudinal dark brown stripes and frons mostly shining black, but medially with a narrow longitudinal yellow marking and a narrow median U-shaped yellow marking at ventral corner ( Fig. 3B-D).

Etymology
The new species is named after its type locality, i.e., Azad Kashmir, Pakistan.  . Vertex moderately raised, dark brown with yellow markings; in frontal view dark brown, dorsally with a pair of C-shaped yellow markings, which are connected to ocular rim; in dorsal view dark brown but medially yellow, with a rounded dark brown marking at middle; epicranial area black, with longitudinal transverse grooves, covered with short brownish pubescence. Frons shining black, but medially with a narrow longitudinal yellow marking and a narrow median U-shaped yellow marking at ventral corner, covered with short brownish pubescence. Occiput and postorbital sclerites yellow. Clypeus pale yellow (Fig. 3B) or medially with two small rounded dark brown spots in some specimens (Fig. 3C), and with four median long brown setae. Labrum brownish yellow, with erected brownish setae at proximal margin. Genae pale yellow. Maxillary and labial palps pale yellow, with terminal labial palpomere spindleshaped, palpimacula brownish, small and circular, with short pale yellow setae. Antennae black, covered with short black setae, dorsal ring of scape and pedicel yellowish brown, fl agellum black with several distal fl agellomeres widened and moderately fl attened. Antennal sclerite yellow (Fig. 3B). THORAX (Fig. 3A). Pronotum slightly wider than long, yellow, with two median longitudinal dark brown stripes; lateral margins narrowly dark brown after anterior transverse furrow in lateral view; covered with long sparse black setae, but anterolateral margins with short, black and pale yellow setae. Mesonotum yellow with a median and lateral dark brown stripe; prescutum laterally with a pair of rounded yellow spots, covered with long sparse black setae; mesoscutum with a median and lateral dark brown stripe, lateral stripes limited to proximal ⅔, covered with short yellow setae; metascutellum yellow with a broad median longitudinal dark brown stripe, covered with long sparse yellow setae. Metanotum yellow with a median and lateral dark brown stripe, lateral stripes limited to proximal half of metascutum, covered with short yellow setae. Pleuron yellow, with a median longitudinal dark brown stripe, covered with short sparse yellow setae (Fig. 3F). LEGS (Fig. 3F). Foreleg: coxa and trochanter yellow, covered with short yellow setae. Femur yellow, covered with short, black and brownish yellow setae, proximal half with a few long black setae; femoral sense hair as long as proximal ½ of profemora. Tibia yellow, with mixed, short and thick black setae,    antennal cleaning setae yellow; tibial spurs brownish, straight, as long as Ta1. Tarsomeres yellow, covered with short black setae; Ta1 equal to combined length of Ta2-Ta3; Ta2, Ta3 and Ta4 nearly equal in size; Ta5 equal to combined length of Ta1-Ta4. Pretarsal claws brownish, curved. Mid leg similar to foreleg. Hind leg similar to mid leg, but femoral sense hair absent. WINGS (Fig. 2). Forewing: relatively broad, subacute at apex; membrane hyaline; longitudinal veins dark brown, except Sc with alternate dark brown and yellow patches, covered with sparse short black setae; costal area with interconnected crossveins proximal to pterostigma for at least ⅓ of forewing length; seven to nine presectoral crossveins; initial branching point of CuA at same level to Rs origin; Rs with 12-16 branches; pterostigma milky white; anterior Banksian line absent; posterior Banksian line present. Hind wing: slightly longer than forewing, acutely pointed at apex; membrane hyaline; longitudinal veins dark brown, except Sc with alternate dark brown and yellow patches, covered with sparse short black setae; costal veinlets simple, except a few marginally forked crossveins around poststigmal area; fi ve presectoral crossveins; median fork proximal to Rs origin; Rs with 14-18 branches; pterostigma milky white, relatively smaller than that on forewing; anterior Banksian line absent; posterior Banksian line present; pilula axillaris with rounded knob, covered with dense brown setae. ABDOMEN ( Fig. 1). Tergites brownish, covered with short brownish yellow setae. Sterites brownish yellow, covered with short brownish yellow setae. MALE GENITALIA (Fig. 4C-G). Tergum 9 trapezoidal, with anterior margin slightly prominent in lateral view. Sternum 9 slenderly triangular in ventral view, covered with long black setae at distal half. Ectoproct nearly rectangular in lateral view, posterodorsal margin rounded, covered with yellow setae, posteroventral corner slightly prominent, covered with long thick black setae. Gonocoxites 11 highly sclerotized, lateral arms straight, apex curved ventrad in dorsal view. Gonostylus 11 slightly prominent in ventral view. Gonocoxites 9 broad and elongate, curved with pointed apex in lateral view. FEMALE GENITALIA (Fig. 4A-B). Tergum 9 quadrate in lateral view, with short black setae. Ectoproct subquadrate, posterodorsal margin rounded, with short yellow setae, but proximal ⅓ with robust digging setae. Anterior gonocoxites 8 short, as long as wide, with thick long black setae. Posterior gonocoxites 8 long, digitiform, with long black setae. Gonocoxites 9 broad and rounded, with robust digging setae, anterolaterally with a bunch of erected short black setae at proximal ⅓.

Diagnosis
Myrmeleon is similar to Baliga by the presence of anterior gonocoxites 8 relatively shorter than posterior gonocoxites 8 in the female genitalia ( Fig. 9A-B) but can be distinguished by the absence of interconnected crossveins in the costal area of forewing (Fig. 7).
Key to species of the genus Myrmeleon from Pakistan* 1. Vertex wholly black, without yellow markings (Fig. 18C)  2. Clypeus yellow with two median rounded brownish markings; pronotum dark brown, medially with a narrow longitudinal complete yellow stripe, rounded in center, laterally with a narrow yellow stripe at proximal half (Hölzel 1972: fi g. 97); male gonocoxite 9 arcuate at distal margin in ventral view (Hölzel 1972
THORAX (Fig. 8A, C). Pronotum slightly wider than long, yellow, with two median longitudinal dark brown stripes, separated by a narrow yellow central stripe, lateral margins yellow, covered with long yellow setae. Mesonotum dark brown, medially with faintly brownish yellow marking, distal margin yellow, covered with sparse yellow setae, but prescutum with long dark brown setae. Metanotum dark brown, medially with faintly brownish yellow markings, distal margin of metascutellum yellow, covered with sparse yellow setae. Pleuron dark brown, covered with sparse yellow setae (Fig. 8E).
LEGS (Fig. 8E). Foreleg: coxa and trochanter yellow, covered with short yellow setae. Femur yellow, posterolaterally brownish at apex, covered with short black setae, but posterolaterally with a few long black setae at proximal ½ and ventrally with short yellow setae; femoral sense hair shorter than proximal ½ of profemora. Tibia yellow, anterolaterally brownish, covered with short black setae, posterolaterally with a few long black setae, antennal cleaning setae yellow; tibial spurs brownish, straight, as long as Ta1.
Tarsomeres yellow, covered with short black setae; Ta1 equal to combined length of Ta2-Ta4; Ta2, Ta3 and Ta4 nearly equal in size; Ta5 equal to combined length of Ta1-Ta3. Pretarsal claws brownish, moderately curved. Mid leg: coxa and trochanter similar to foreleg. Femur yellow, anterolaterally brownish, covered with short black setae, ventrally with soft yellow and laterally with a few long black setae at proximal half; femoral sense hair shorter longer than proximal half of mid femora. Tibia similar to foreleg, but laterally     with a few long black setae; tibial spurs similar to foreleg. Tarsomeres and pretarsal claws similar to foreleg. Hind leg: coxa and trochanter similar to mid leg. Femur yellow, with distinct subapical brownish ring, covered with short black setae, but at proximal half with a few long black setae; femoral sense hair absent. Tibia yellow, ventrally brownish, covered with short black setae, ventrally with a row of long black setae; tibial spurs similar to middle leg. Tarsomeres and pretarsal claws similar to mid leg. WINGS (Fig. 7). Forewing: slightly longer and wider than hind wing, subacute at apex; membrane hyaline; costal area slightly narrow at proximal region; venation yellow, except Sc with alternate brownish and yellow patches at proximal half, covered with sparse short black setae; poststigmal area with a few interconnected crossveins; seven to nine presectoral crossveins; initial branching point of CuA at same level or proximal to Rs origin; Rs with 10-13 branches; CuP origin at same level to basal crossveins, fused with 1A after a short free base; pterostigma small, milky white; anterior Banksian line absent; posterior Banksian line present. Hind wing: relatively narrower than forewing, acute at apex; membrane hyaline; venation similar to forewing; four presectoral crossveins; median fork proximal to Rs origin; Rs with 12-14 branches; pterostigma indistinct; anterior Banksian line absent; posterior Banksian line present; pilula axillaris small, with rounded knob, covered with dense brown setae. ABDOMEN (Fig. 6). Tergites dark brown, but in some specimen with a narrow transverse yellow stripe at distal margin of terga 4-8, covered with short brownish yellow setae. Sternites dark brown, covered with short brownish yellow setae.
MALE GENITALIA (Fig. 9C-G). Tergum 9 trapezoidal, with anterior margin slightly prominent in lateral view. Sternum 9 ovoid in ventral view, covered with elongated black setae at distal half. Ectoproct nearly rectangular in lateral view, covered with yellow setae at distal half, posteroventral corner at proximal half with long thick black setae in lateral view. Gonocoxites 11 highly sclerotized, lateral arms straight, posterolaterally wide and rounded in dorsal view. Gonostylus 11 slightly prominent in dorsal view. Gonocoxites 9 broad and elongated, proximally diverged in dorsal view, distally rounded in lateral view.
FEMALE GENITALIA (Fig. 9A-B). Tergum 9 quadrate in lateral view, covered with short black setae. Ectoproct rounded in lateral view, posterodorsal margin rounded, with short yellow setae, but proximal ⅓ with robust digging setae. Anterior gonocoxites 8 short, as long as wide, covered with thick long black setae. Posterior gonocoxites 8 long, digitiform, covered with long black setae. Gonocoxites 9 broad and rounded, covered with robust digging setae, anterolaterally with a bunch of erected short black setae at proximal ⅓. Pregenital plate small, pointed at apex in ventral view.

Note
Myrmeleon tenuipennis is rarely mentioned in literature since its original description (Rambur 1842;Ghosh 1983;Stange 2004). Previously, it was only known from India, Sri Lanka, and Vietnam (Stange 2004;Oswald 2020). But we thought this is the most common antlion species in Pakistan, previously misidentifi ed as M. assamensis. The marking patterns on frons and pronotum of M. assamensis reported from Pakistan : fi g. 1a) and the paratype of Myrmeleon bimaculatus Yang, 1999 (Fig. 11) from China are almost identical to the type photographs of M. fryeri Navás, 1914 (Fig. 10). However, it can be distinguished from these closely related species by the presence of two yellow markings on vertex in dorsal view (with four yellow markings in M. assamensis: two at median and two at lateral margins in dorsal view). After careful examination of the holotype photographs of M. fryeri (Fig. 10), which is a junior synonym of M. tenuipennis proposed by Esben-Petersen (1931), it is concluded that the specimens presently collected from Pakistan are M. tenuipennis based on the presence of two yellow markings on vertex, instead of four in M. assamensis. We also examined the paratype of M. bimaculatus Yang, 1999 (holotype lost), and confi rm that this species is a synonym of M. tenuipennis. Myrmeleon tenuipennis is widely distributed in coastal areas of southern China.

Diagnosis
It can be distinguished by the distinctive head and thoracic markings: frons black, except for ventral corner yellow (Fig. 14B); vertex black, median and posterior portions with yellow markings in dorsal view; pronotum yellow, medially with a longitudinal brownish stripe, anterior transverse furrow dark brown, distally with a pair of well-separated transverse brownish stripes (Fig. 14C); wings narrowly elongated, acutely pointed at apex, initial branching point of CuA distad Rs origin (Fig. 13). Moreover, male genitalia is distinctive among species of Myrmeleon in Pakistan: gonocoxites 11 highly sclerotized, lateral arms elongated; gonostylus 11 rounded in lateral view; gonocoxites 9 narrow and elongated, wider in lateral view with pointed apex.

Re-description
MEASUREMENT (♂n = 1). Forewing: length 26.2 mm, width 5.5 mm; hind wing: length 26.2 mm, width 4.6 mm; body length: 24.0 mm. HEAD ( Fig. 14B-C). Vertex moderately raised; in frontal view black, without yellow markings; in dorsal view black, medially with a pair of transverse and posteriorly with a pair of longitudinal yellow markings; epicranial area black, with longitudinal grooves, covered with short brownish pubescence. Frons black, but ventral corner yellow, covered with short yellowish pubescence. Occiput and postorbital sclerites yellow. Clypeus yellow, medially with two indistinct dark brown markings. Labrum yellow, covered with erected brownish setae at proximal margin. Genae pale yellow. Maxillary and labial palps pale yellow, terminal labial palpomere spindle-shaped, palpimacula brownish, small and circular, with short black setae. Antennae brownish, dorsal ring of scape and pedicel yellow, covered with short black setae, fl agellum brownish with proximal and distal margin dark brown. Antennal sclerites yellow (Fig. 14B).
THORAX (Fig. 14A). Pronotum slightly wider than long, yellow, medially with a longitudinal brownish stripe, slightly interrupted at anterior transverse furrow; dark brown stripe along anterior transverse furrow, not reaching at lateral margins; distally with a pair of well-separated transverse brownish stripes; covered with sparse yellow setae. Mesonotum dark brown; prescutum laterally with a narrow yellow stripe; mesoscutum with yellow markings on median and posterolateral margins; mesoscutellum at distal ⅔ yellow; covered with sparse yellow setae, but prescutum with long dark brown setae. Metanotum dark brown; prescutum medially with faintly brownish longitudinal yellow marking; metascutum distally with a pair of large yellow markings; metascutellum laterally and distally brownish yellow; covered with sparse yellow setae. Pleuron dark brown, with yellow markings, covered with sparse yellow setae (Fig. 14E).
LEGS (Fig. 14E). Foreleg: coxa and trochanter yellow, with short yellow setae. Femur yellow, light brownish at distal ½, with short black setae, but posterolaterally with a few long black setae at proximal half; femoral sense hair shorter than proximal ½ of profemora. Tibia yellow, with mixed, short and long black setae at proximal half, antennal cleaning setae yellow; tibial spurs brownish, straight, as long as Ta1. Tarsomeres yellow, with short black setae; Ta1 equal to combined length of Ta2-Ta3; Ta2, Ta3 and Ta4 nearly equal in size; Ta5 equal to combined length of Ta1-Ta3. Pretarsal claws brownish, curved. Mid leg: coxa yellow, slightly brownish at proximal ½, with yellow setae. Trochanter yellow, with short black setae. Femur yellow, light brownish at distal ½, covered with short black setae, laterally with a few long black setae at proximal half; femoral sense hair shorter than proximal half of mid femora. Tibia yellow, covered with mixed, short and long black setae; tibial spurs similar to foreleg. Tarsomeres and pretarsal claws similar to foreleg. Hind leg: coxa and trochanter similar to mid leg. Femur yellow, light brownish at distal ⅓, covered with short black setae, proximal half with a few long black setae; femoral sense hair absent. Tibia, tarsomeres, and pretarsal claws are similar to mid leg. WINGS (Fig. 13). Forewing as long as hind wing, acute at apex; membrane hyaline; costal area slightly narrow at proximal region; venation brownish yellow, covered with sparse short black setae; poststigmal present. Hind wing: relatively narrow, acute at apex; membrane hyaline; venation similar to forewing; fi ve presectoral crossveins; median fork at proximal to Rs origin; Rs with 12 branches; pterostigma indistinct; anterior Banksian line absent; posterior Banksian indistinct; pilula axillaris small, with rounded knob covered with dense brown setae. ABDOMEN (Fig. 12). Tergites dark brown, distally with a narrow yellow stripe on terga 1-8, lateral margins yellow, covered with short brownish yellow setae. Sternites dark brown, distally with a narrow yellowish stripe, covered with short brownish yellow setae.
MALE GENITALIA (Fig. 15). Tergum 9 subtrapezoidal in lateral view. Sternum 9 ovate-shaped, covered with long black setae at distal ⅓. Ectoproct nearly rectangular in lateral view, posteroventral corner slightly prominent, covered with long thick black setae. Gonocoxites 11 highly sclerotized, lateral arms elongated, apex broad and curved ventrad in dorsal view. Gonostylus 11 rounded in lateral view. Gonocoxites 9 narrow and elongated, wider in lateral view with pointed apex.  (Fig. 21). In conclusion, a combined morphological and molecular data should be applied for all subspecies to resolve the status of these geographically isolated subspecies.

Diagnosis
Body coloration generally dark brown; frons and vertex wholly black, without yellow markings ( Fig. 18B-C); pronotum slightly wider than long, mostly yellow, with two broad median dark brown stripes, narrowly separated by a central yellow line; meso-and metanotum dark brown, posteriorly yellow, covered with scattered fi ne yellowish setae (Fig. 18A).
THORAX (Fig. 18A). Pronotum slightly wider than long, yellow, medially with pair of longitudinal dark brown stripes, separated by a narrow central yellow line, covered with short sparse yellow setae, anterolaterally with short black setae, posterolaterally and distally with a few long black setae. Mesonotum dark brown, covered with sparse yellow setae; posterolateral margins on pre-and mesoscutum with yellow markings, prescutum covered with long dark brown setae; metascutellum distally with a narrow yellow stripe, covered with sparse yellow setae. Metanotum dark brown, metascutellum distally with a narrow yellow stripe, covered with sparse yellow setae. Pleuron dark brown, covered with sparse yellow setae (Fig. 18E).
LEGS (Fig. 18E). Foreleg: coxa yellow, slightly brownish at proximal ⅓, with short yellow setae. Trochanter yellow, covered with mixed, short, black and yellow setae. Femur yellow, posterolaterally brownish at apex, covered with short black setae, proximal half with a few long black setae, proximal ⅓ with short yellow setae; femoral sense hair shorter than proximal half of profemora. Tibia yellow, laterally brownish in some specimens, distally shiny black, with short black setae, posterolaterally with a few long black setae, antennal cleaning setae yellow; tibial spurs brownish, straight, as long as Ta1. Tarsomeres yellow, distally each tarsomere dark brown, with short black setae; Ta1 longer than Ta2; Ta2, Ta3 and Ta4 nearly equal in size; Ta5 nearly equal to combined length of Ta1-Ta4. Pretarsal claws brownish, moderately curved. Mid leg: coxa and trochanter similar to foreleg. Femur yellow, anterolaterally dark brown, covered with short black setae, proximal ⅓ with short yellow setae but proximal half with a few long black setae; femoral sense hair shorter than proximal half of mid femora. Tibia yellow, anterolaterally brownish, distally shiny black, covered with short and long black setae; tibial spurs, tarsomeres and pretarsal claws similar to foreleg. Hind leg: coxa and trochanter similar to mid leg. Femur yellow, anterolaterally at distal ⅓ brownish, covered with long black setae, but proximal half with short yellow setae; femoral sense hair absent. Tibia, tibial spurs, tarsomeres, and pretarsal claws similar to mid leg. WINGS (Fig. 17). Forewing: relatively broad, slightly longer than hind wing, subacute at apex; membrane hyaline; costal area slightly narrow at proximal region; longitudinal veins yellow, except Sc and Cu at proximal half with alternate dark brown and yellow patches; crossveins yellow, except cubital area after posterior Banksian line with crossveins black; six to nine presectoral crossveins; initial branching point of CuA proximal to Rs origin; Rs with 11-14 branches; pterostigma small, milky white; anterior Banksian line indistinct as compared to more prominent posterior Banksian line which is proximally brownish black. Hind wing: relatively narrower than forewing, acute at apex; membrane hyaline; longitudinal veins yellow except Sc at proximal half with alternate dark brown and yellow patches; crossveins yellow, but median area after posterior Banksian line with crossveins black; fi ve to six presectoral crossveins; median fork proximal to Rs origin; Rs with 12-14 branches; pterostigma milky white; anterior Banksian line absent; posterior Banksian line indistinct; pilula axillaris with rounded knob, covered with dense brown setae. ABDOMEN (Fig. 16). Tergites dark brown, distally with a narrow yellow stripe on terga 4-8; terga 4-7 relatively broader in both sexes; covered with short yellowish setae, but posterior and posterolateral margins of terga 6-7 with mixed, short, black and brownish setae, tergum 8 with short black setae. Sternites dark brown, sterna 7-8 each with distally a narrow yellow stripe; covered with short yellowish setae.
MALE GENITALIA (Fig. 19C-G). Tergum 9 trapezoidal, with anterior margin slightly prominent in middle in lateral view. Sternum 9 triangular in ventral view, covered with long and elongated black setae. Ectoproct nearly rectangular in lateral view, posterodorsal margin rounded, covered with yellow setae, posteroventral corner slightly prominent, covered with long thick black setae. Gonocoxites 11 highly sclerotized, lateral arms straight, rounded at apex in ventral view, apex curved ventrad in dorsal view. Gonostylus 11 cone-shaped, prominent in ventral view. Gonocoxites 9 separated, narrow and straight proximally, wide and curved distally with pointed apex in lateral view.

Diagnosis
Myrmeleon assamensis can be distinguished by the marking patterns on vertex and pronotum: vertex black, posteriorly with four yellow markings, two at middle and two at lateral margins in dorsal view; pronotum yellow, medially with a pair of dark brown stripes, which are separated by a narrow median yellow line.

Note
In the original description (solely based on male), Ghosh (1984) characterized this species based on the presence of two black stripes on the pronotum and the number of presectoral crossveins and the radial branches in both wings. Subsequently, Ghosh (1990) described the female of this species and found that the number of presectoral crossveins and the radial branches are different from that in the male previously described by him. Later, Iqbal & Yousuf (1992: fi g. 2) re-described this species based on specimens from Pakistan and provided the line drawings of the head and pronotum in dorsal view as well as the male genitalia. Recently, Akhtar et al. (2018: fi g. 1a) also recorded this species in Pakistan, which, however, is a case of misidentifi cation of M. tenuipennis (see Note to this species). In the original description of Myrmeleon assamensis, the marking patterns on vertex are as follows: vertex at distally with two longitudinal and two transverse dark spots in dorsal view (vertex with only two longitudinal yellow markings in M. tenuipennis). No new specimens of this species were found in this study.

Diagnosis
Myrmeleon bore can be characterized by wholly dark brown vertex without yellow markings and the pronotum with lateral margins narrowly yellow on proximal half (Aspöck et al. 1980 (Akhtar et al. 2018: fi g. 2a). This character suggests that the specimens from Pakistan identifi ed as Myrmelon bore belong to a different species.

Remarks
Myrmeleon bore seems widely distributed in the Palaearctic Region and was recently recorded from Pakistan. Notably, Enza otiosus Navás, 1912 has long been considered a secondary synonym of M. bore (Stange 2004;Sekimoto 2014;Wang et al. 2018

Diagnosis
Myrmeleon clothilde can be characterized by wholly dark brown vertex with yellow markings at distal half in dorsal view; pronotum dark brown, laterally slightly narrow yellow at proximal half; medially with two narrow longitudinal yellow stripes at proximal to anterior transverse furrow (Iqbal & Yousuf 1992: fi g. 1a).

Note
Since its original description, this species was rarely mentioned in literature (Iqbal & Yousuf 1992, 1997Ghosh 2000;Stange 2004). After reviewing the aforementioned literature from Pakistan and India, we found that the male genitalia of this species have not been described so far. However, further additional data on the male genitalia and distribution of this rarely known species in Pakistan need to be updated in further studies. No new specimens of this species were found in this study.

Diagnosis
Myrmeleon noacki can be characterized by a wholly dark brown vertex, without yellow markings; pronotum dark brown, laterally yellow, medially with a narrow longitudinal yellow line at proximal half wings lack pipula axillaris in males; apex of male gonocoxites 9 narrowly arcuated at anterolateral margins in ventral view.

Note
Myrmeleon noacki is known from the southeastern parts of Europe to Turkey, and was recently reported from Iran and Pakistan Hajiesmaeilian et al. 2020). The reports of Myrmelon noacki from Pakistan need to be re-evaluated and compared with European specimens. Typically, this species is characterized by a narrow median longitudinal yellow marking at proximal half of pronotum and the male genitalia with gonocoxites 9 at anterolateral margins arcuated at anterolateral margins at apex in ventral view (see Ohm 1965: fi gs 2, 6; Hajiesmaeilian et al. 2020: fi gs 10, 14). The prothoracic markings and the shape of male genitalia of this species recorded from Pakistan match the typical diagnosis for M. paghmanus: pronotum dark brown, medially with a narrow longitudinal yellow marking, rounded at middle; male gonocoxites 9 arcuate at distal margin in ventral view (see Akhtar et al. 2018: fi g. 5a-b; Hölzel 1972: fi gs 97, 101-102). No new specimens of this species were found in this study.

Diagnosis
Myrmeleon paghmanus can be characterized by the combination of the following characters: clypeus yellow, medially with two rounded brownish markings; pronotum dark brown, medially with a narrow longitudinal complete yellow stripe, laterally with a narrow yellow stripe at proximal half wings lack pipula axillaris in males; male gonocoxites 9 arcuate at distal margin in ventral view.

Note
Up till now, this species is only known from Afghanistan and Pakistan. No new specimens of this species were found in this study.

Molecular identifi cation
The present phylogenetic analysis based on COI and 16S rRNA genes shows that there is strong support for the monophyly of Baliga clade for Japanese species by BI, ML, and NJ methods, which, however, did not comprise Baliga kashmirensis sp. nov. from Pakistan. Based on COI genes, Baliga kashmirensis sp. nov. was assigned to be within a monophylum with M. tenuipennis and M. taiwanensis Miller & Stange, 1999. However, the monophyly of Baliga kashmirensis sp. nov. with M. tenuipennis, M. hyalinus, M. trivialis, and the M. formicarius clade is recovered with relatively low nodal support values. For now, the present phylogenetic analysis is largely focused on species identifi cation due to incomplete taxon sampling. The genetic divergence between B. kashmirensis sp. nov. and the species of Myrmeleon was 0.139-0.188 and that between this new species and the other species of Baliga was 0.153-0.186. The greatest intraspecifi c divergence (0.049) was found respectively in B. ryukyuensis Hayashi & Matsumoto, 2020 and M. hyalinus. The minimum and maximum interspecifi c genetic divergence between species of Baliga and Myrmeleon ranged from 0.074-0.186, and 0.123-0.188, respectively.

Discussion
In most recent taxonomic and molecular studies on Myrmeleontini from Japan, Hayashi et al. (2020) considered Baliga as a valid genus based on the generic classifi cation system proposed by Stange (2004). According to Stange (2004), Baliga can be characterized by the presence of interconnected crossveins proximal to pterostigma in the forewing and the anterior gonocoxites 8 shorter than the posterior gonocoxites 8 in the female genitalia. However, the BI, ML and NJ trees herein reconstructed respectively based on COI and 16S rRNA genes for six species of Baliga and 11 Myrmeleon (Table 1; Fig. 21) albeit receiving low supports at deep-level nodes recovered Myrmeleon as paraphyletic. Similar results were also recovered in the recent molecular phylogenetic studies by Michel et al. (2017) and Machado et al. (2019). In Hayashi et al. (2020), the species of Baliga and those of Myrmeleon, respectively, constituted a monophylum, which, however, might be due to incomplete taxon sampling. Nevertheless, here we do not propose any new generic synonym but still follow the generic classifi cation system of Stange (2004). The validity of Baliga needs a major phylogenetic revision with larger datasets in future studies. Furthermore, the additional proposed synonymized genera by Machado et al. (2019) need to be deciphered in future studies. In addition, concerning the great intraspecifi c divergence, it is necessary to clarify the taxonomic status of all subspecies of M. hyalinus and the two insular populations of B. ryukyuensis (Amami/Tokunoshima and Okinawa) based on both morphological and molecular data.
With respect to the distribution of antlions in Pakistan, our results corroborate the mixed fauna from the Oriental and Palaearctic regions in the northern parts of Pakistan (Fig. 20) due to the unique geographical position -the extreme edge of western Himalayas and the junction point of the world's two largest zoogeographical regions: the Oriental and the Palaearctic. The present diversity and distribution of antlions in Pakistan is consistent with our recent studies on the subfamily Ascalaphinae (Neuroptera: Myrmeleontidae) and the following genera of Myrmeleontidae and Megaloptera from northern Pakistan: Distoleon Banks, 1910 (Neuroptera: Myrmeleontidae), Nevromus Rambur, 1842 and Protohermes van der Weele, 1907 (Megaloptera: Corydalidae) (Hassan et al. 2019(Hassan et al. , 2020a(Hassan et al. , 2020bHassan & Liu 2021). Nevertheless, this pattern may be also due to the present limitation of sampling (extensive collecting primarily confi ned to the northern parts of Pakistan). Broader sampling across the country, particularly for the southern parts, may reveal the true diversity and distribution of antlions in the future.