New Cretaceous antlion-like lacewings promote a phylogenetic reappraisal of the extinct myrmeleontoid family Babinskaiidae

Babinskaiidae is an extinct family of the lacewing superfamily Myrmeleontoidea, currently only recorded from the Cretaceous. The phylogenetic position of this family is elusive, with inconsistent inferences in previous studies. Here we report on three new genera and species of Babinskaiidae from the mid-Cretaceous Kachin amber of Myanmar, namely Calobabinskaia xiai gen. et sp. nov., Stenobabinskaia punctata gen. et sp. nov., and Xiaobabinskaia lepidotricha gen. et sp. nov. These new babinskaiids are featured by having specialized characters, such as the rich number of presectoral crossveins and the presence of scaly setae on forewing costal vein, which have not yet been found in this family. The exquisite preservation of the Kachin amber babinskaiids facilitate a reappraisal of the phylogenetic placement of this family based on adult morphological characters. Our result from the phylogenetic inference combining the data from fossil and extant myrmeleontoids recovered a monophyletic clade composed of Babinskaiidae and another extinct family Cratosmylidae, and further assigned this clade to be sister group to a clade including Nemopteridae, Palaeoleontidae, and Myrmeleontidae. Babinskaiidae appears to be a transitional lineage between Nymphidae and advanced myrmeleontoids, with ancient morphological diversification.

www.nature.com/scientificreports/ from R slightly distal to midpoint of wing; RP pectinately branched from its proximal 1/3, with one rp-ma crossvein in left forewing and three in right forewing between stem of RP and MA; RP with eight branches, mostly simple, but RP4 deeply forked; RP1 fused with RP2 distally, RP3 fused with anterior branch of RP4 distally, RP7 fused with RP8 distally; one gradate series of four crossveins present near branching points of RP branches; MA simple; MP long and nearly straight, pectinately branched from distal 1/4, with 18 branches, mostly bearing a marginal fork; 27 crossveins present between MP and CuA; CuA and CuP diverging near wing base; CuA pectinately branched, with six branches, mostly bearing a marginal fork; CuP long, proximally fused with A1, terminating at same level of diverging point between RP and MA, with 28 pectinate, short and simple branches; 29 cua-cup crossveins present; A2 and A3 simple, connected with each other by a short a2-a3 crossvein. Hind wing: Proximally distinctly narrowed, remaining part nearly as wide as forewing; costal space slightly wider than subcostal space, with at least 24 simple crossveins on proximal 4/5, and 25 crossveins and veinlets of ScP + RA on distal 1/5; subcostal crossveins absent; 19 presectoral crossveins present; RP + MA originating distal to midpoint of wing; two rp-ma crossveins present between stem of RP and MA; RP pectinately branched from its proximal 1/3 into eight simple branches, with distal-most two ones fused distally; one gradate series of four crossveins present near branching points of RP branches; MP1 and MP2 straight and long, both pectinately branched from distal 1/5, respectively with 15 and seven branches, mostly bearing a marginal fork; CuA long, terminating at same level of diverging point between MA and RP, with ca. 25 pectinate simple branches.
Etymology The new species is dedicated to Mr. Fangyuan Xia, who kindly offered the specimen (the holotype) of this new species for present study. Prothorax much narrower than head, slightly wider than long; meso-and metathorax slightly wider than prothorax. Wings narrowly elongate, ca. 6.5 times as long as wide; wing spots present on apex of both fore-and hind wings, mostly on some crossveins of branching region of RP; single trichosor present between veins along distal margin in both fore-and hind wings.
Forewing: Costal space strongly narrowed, slightly wider than subcostal space, but narrower than radial space, with ca. 30 simple crossveins on proximal 3/4, and ca. 45 crossveins and veinlets of ScP + RA on distal 1/4, mostly forked marginally; subcostal crossveins absent; 11 presectoral crossveins present, one of them sigmoid and acutely angled with R; RP + MA originated from R slightly proximal to midpoint of wing; RP pectinately branched from its proximal 1/5, with one rp-ma crossvein between stem of RP and MA; RP with 13 branches, mostly simple, but RP5 and RP6 deeply forked; RP8 and RP9, RP12 and RP13 fused distally; at least four gradate series of crossveins present among branches of RP and MA at proximal half; an additional distal gradate series of crossveins present; MA only with a marginal fork; MP long and nearly straight, pectinately branched from distal 1/4, with 17 branches, mostly bearing a marginal fork; 22 crossveins present between MP and CuA; CuA and CuP diverging near wing base; CuA pectinately branched, with at least 10 branches, most of which are simple; CuP long, proximally fused with A1, terminating at same level of diverging point between RP and MA, pectinately branched with at least 23 short and simple branches; 18 cua-cup crossveins present; A2 and A3 simple, connected with each other by a short a2-a3 crossvein.
Hind wing: Proximally distinctly narrowed, remaining part nearly as wide as forewing; costal space narrow, nearly twice as wide as subcostal space, with at least 28 crossveins on proximal 3/4, and 45 crossveins and veinlets of ScP + RA on distal 1/4, mostly forked marginally; subcostal crossveins absent; at least 10 presectoral crossveins present, one of them sigmoid and acutely angled with R; RP + MA originating slightly proximal to midpoint of  Abdomen slenderly elongate, with segments 4-5 slightly widened. Male genitalia: Tergum 9 slightly shorter than tergum 8; sternum 9 large, about twice as long as tergum 9 plus ectoproct, straightly extending posteriad; putative gonocoxite 9 present, but probably detached from its original position, strongly curved medially with acute tip; putative gonostyli 11 present as a pair of long spinous sclerites, which are slightly shorter than sternum 9; ectoprocts paired, subtrapezoidal in lateral view, with large ovoid callus cerci, and distally with some short teeth.
Etymology The specific epithet "punctata" refers to the occurrence of wing spots in the new species.  Etymology From "Xiao-" and "Babinskaia" (the type genus-group name of Babinskaiidae) in reference to Mrs. Xiao Jia, who kindly offered the specimen (the holotype) of the genus type for our study. Gender: Feminine.   Head with vertex medially domed; compound eyes large, semi-globular; antenna partly preserved; scape much wider and slightly longer than pedicel; flagellum with 22 flagellomeres preserved, each flagellomere distinctly shorter than and almost as wide as scape.
Prothorax narrower than head, slightly longer than wide; meso-and metathorax robust, much wider than prothorax. Wings broad, ca. 3.5 times as long as wide, transparent and immaculate; single trichosor between veins along distal margin but 3-5 trichosors present between distal costal crossveins and branches of CuA and CuP in forewing, or branches of MP2 and CuA in hind wing.
Forewing: Wing base strongly narrowed, with a row of scaly setae on proximal section of costal vein, and a number of thick setae along proximal sections of ScP, MP, and A1; costal space gradually widened distally, nearly three times as wide as subcostal space, distally distinctly wider than radial space, with ca. 27 simple crossveins on proximal 2/3, and 21 crossveins and veinlets of ScP + RA on distal 1/3, mostly marginally forked; subcostal crossveins absent; six presectoral crossveins present; RP + MA originating at proximal 1/3 of wing; RP pectinately branched from its proximal 1/5, with one rp-ma crossvein between stem of RP and MA; RP with eight branches, mostly bearing a marginal fork; 16 crossveins present among branches of RP and MA on proximal half; a distal gradate series of crossveins present; MA slightly zig-zagged, with a marginal fork; MP long and straight, branched from distal 1/6, with six branches, mostly deeply forked; 17 crossveins present between MP and CuA; CuA slightly zig-zagged, pectinately branched at distal 1/3, with 10 simple branches; CuP and A1 proximally fused, distally zigzagged, terminating slightly distal to diverging point between RP and MA, pectinately branched into 14 simple branches; A2 approximating stem of A1, distally abruptly curved posteriad; A3 simple, with one a2-a3 crossvein.
Hind wing: Slightly longer and narrower than forewing, with base distinctly narrowed, and with apex acutely pointed and slightly bended posteriad; costal space nearly twice as wide as subcostal space, with 22 mostly simple crossveins on proximal 3/4 and 16 crossveins and veinlets of ScP + RA on distal 1/4, mostly forked marginally; subcostal crossveins absent; five presectoral crossveins present; RP + MA originating at proximal 1/3 of wing; RP pectinately branched from its proximal 1/5 into eight branches, mostly bearing a marginal fork; one rp-ma Legs slender, with sparse short setae; a pair of tibial spurs present; tarsus 5-segmented; tarsomere 1 longest, nearly twice as long as tarsomere 2; tarsomeres 3-5 shortest, nearly half length of tarsomere 2; each tarsomere slightly widened distally; pretarsal claws slender and slightly curved, equal in length and shape; arolium not discernible.
Abdomen slenderly elongate, with segments 4-6 much broader. Female genitalia: Tergum 8 nearly as long as tergum 9 plus ectoproct; ventral sclerites of segment 8 not discernible; tergum 9 nearly rectangular in lateral view; a pair of broad valvate gonocoxites 9 present, proximally with a pair of short and flat lobes, which are rounded at tip; ectoprocts paired and broad, nearly rectangular in lateral view; callus cerci not discernible.
Etymology The specific epithet "lepidotricha" refers to the presence of a row of scaly setae on the proximal section of forewing costal vein in the new species.  Figs. 10, 11 and S1. The presently recovered monophyletic Myrmeleontoidea is supported by the absence of forewing nygmata (character 5:1) and the forewing ScP and RA terminating at or posteriad wing apex (character 13:1). Nymphidae is recovered to be the sister group of the clade including the remaining genera, and its autapomorphies comprise the prothorax slightly elongated anteriad procoxae (3:1), the presence of forewing thrydiate crossveins (character 14:1) and the bifid arolium (character 50:1). The monophyly of the latter clade is supported by the forewing RP + MA diverging slightly distal to wing base (16:1), and the forewing CuA branched near midpoint of wing with 10 or more branches (characters 28:2 and 29:1). Cratosmylidae and Babinskaiidae are clustered together, being sister to the lineage of Nemopteridae + (Myrmeleontidae + Palaeoleontidae). The monophyly of Cratosmylidae + Babinskaii- Within Babinskaiidae, Stenobabinskaia gen. nov. and Calobabinskaia gen. nov. are clustered together based on the presence of more than 10 presectoral crossveins in the forewing (character 15:3), the presence of trapezoidal forewing prehypostigmal cell (character 17:1), the forewing with distance between diverging point respectively of MA and RP1 twice as long as distance between diverging points respectively of RP1 and RP2 (22:1), the forewing MP1 terminating near wing apex (character 25:1), the forewing CuA branches marginally forked (character 31:1), and the extremely long forewing CuP (character 36:3). The remaining babinskaiid genera besides Xiaobabinskaia gen. nov. constitute another monophyletic group supported by the less crossveins in radial space (character 21:1) and the forewing CuP not fused with A1 (38:0), and together sister to Xiaobabinskaia gen. nov. by the zigzagged forewing CuP (37:1). The monophyly of Nemopteridae + (Palaeoleontidae + Myrmeleontidae) is supported by the absence of trichosors (character 4:1), the single forewing MP (24:2), the presence of forewing oblique vein (character 27:1), and the subtriangular bra nching area of MP2 + CuA (character 30:1). The autapomorphies of Nemopteridae comprise the presence of prolonged rostrum (character 2:1), the closely spaced forewing CuA and CuP (character 34:1), and the strongly narrowed and elongated hind wing (characters 41:2 and 42:2). The monophylum including the genera of Myrmeleontidae and Palaeoleontidae is recovered based on the presence of short forewing CuA2 (character 32:1) and short hind wing CuP (character 36:0). The monophyly of Palaeoleontidae is supported based on the forewing RP + MA diverging from a positon near wing base (16:0), the short forewing hypostigmal cell (character 18:0), the forewing radial space with a Banksian line (character 20:1) and the pectinately branched forewing CuA2 with 3-4 branches (33:1). The monophyly of Myrmeleontidae is supported by the strongly dilated antennae (character 1:2), the origin of RP + MA slightly distal to wing base (character 45:1), and the presence of prolonged tibial spur (character 49:1). Within Myrmeleontidae, the extant three subfamilies form a monophyletic clade, being sister to another clade including Pseudonymphinae and Araripeneurinae. Araripeneurinae is nested in a clade including some Cretaceous antlion genera, i.e., Choromyrmeleon Ren and Guo 16 , Nanoleon Hu, Lu and Liu in Lu et al. 6 and Burmaneura Huang et al. 17 .
The overall nodal supports are not strong if considering the Bremer support values/Bootstrap values, which is probably due to the large proportion of missing characters in the fossil taxa. However, weak support does not   www.nature.com/scientificreports/ three apomorphic characters of this family: (1) the forewing MP2 + CuA1 (i.e., CuA in this paper) reaching the apical margin, (2) the origin of forewing RP (i.e., RP + MA in this paper) far from wing base, and (3) the zigzagged forewing CuP. Makarkin et al. (2017) refuted the above characters 1 and 3 to be apomorphic, and outlined four apomorphic characters, i.e., the origin of forewing RP far from wing base (the above character 2), the presence of presectoral crossveins, the single forewing MP, and the reduction of hind wing A2 and A3. Among these characters, only the last character is considered to be the autapomorphy of Babinskaiidae 7 . In Makarkin and Staniczek 10 , however, this autapomorphy was modified to be the very small anal space and the reduction of A3 in the hind wing because distinct hind wing A2 is found in Parababinskaia makarkini Hu et al. 8 and Electrobabinskaia burmana Lu et al. 6,8. In the present result of phylogenetic analysis, some putative autapomorphies of Babinskaiidae mentioned above (e.g., the presence of more than two presectoral crossveins in both fore-and hind wings and the reduced hind wing A2 and A3) turned to be the autapomorphies of Cratosmylidae + Babinskaiidae. Cratosmylidae is composed of two genera from the Lower Cretaceous of Brazil, i.e., Araripenymphes Menon et al. 18 and Cratosmylus Myskowiak et al. 7,19 . The former taxon is placed within Nymphidae 18,19 . This latter taxon was originally established as a subfamily of Osmylidae 19 , but later transferred to Nymphidae by Winterton et al. 20 . The two genera were considered to represent a separate family Cratosmylidae by Makarkin et al. 7 . Cratosmylidae appears to be a transitional lineage between Nymphidae and Babinskaiidae because it has an intermediate type of characters (Fig. 11). For example, Cratosmylidae and Nymphidae have a more or less proximal position of the forewing RP + MA origin and a deeply branched forewing MP, while Cratosmylidae and Babinskaiidae have several presectoral crossveins. Our result supports that Araripenymphes and Cratosmylus should not belong to Nymphidae (Fig. 11, S1). The closer relationship between Cratosmylidae and Babinskaiidae, as well as the paraphyly of the former family herein recovered, invokes another option that these two taxa may constitute a single family, with Cratosmylidae treated to be a subfamily of Babinskaiidae, as they share some similar apomorphic characters. Nevertheless, due to the scarcity of the cratosmylid fossils, we here retain the current classification until more materials are available for further evaluation.
Phylogenetic position of Babinskaiidae. In the phylogenetic analysis of Myrmeleontoidea combining fossil and extant families 13 , Babinskaiidae was recovered to be the sister group of the clade including most myrmeleontoid families except Nymphidae, which is supported by the presence of forewing presectoral crossveins and forewing MP2 fused with CuA1. However, the forewing presectoral crossveins is absent in many genera of the families/subfamilies within the above clade, such as Araripeneurinae, Pseudonymphinae, Palaeoleontidae, etc. Besides, no oblique vein that is indicative of the fusion between forewing MP2 and CuA1 is found in any babinskaiids 6,7,21,22 . Our result is generally consistent with that in Martins-Neto 13 concerning the position of Babinskaiidae + Cratosmylidae. However, the sister-group relationship between Babinskaiidae + Cratosmylidae and Nemopteridae + (Myrmeleontidae + Palaeoleontidae) is herein supported by the forewing RP + MA diverging from a position slightly distal to wing base, the forewing CuA initially branched near the midpoint of wing, and the forewing CuA with 10 or more branches. Nonetheless, this argument is also not strong enough because the second character state is also present in some genera of Nymphidae and the third character state is absent in most antlions. Given the complex and sometimes unpredictable evolutionary pattern of insect wing venations 23 , the presently used wing characters may not provide sufficient phylogenetic signal to resolve the higher phylogeny of Myrmeleontoidea, especially including many fossil taxa. However, the morphological arguments in Makarkin et al. 7 placing Babinskaiidae together with Nymphidae in Nymphidoidae, i.e., the presence of trichosors and the completely separated forewing MP and CuA, are obviously attributed to the plesiomorphic condition in Neuroptera, which was also mentioned by these authors. Therefore, the phylogenetic position of Babinskaiidae herein recovered is still preferable, but awaits further evidence for corroboration.
Besides the adult characters, the larval morphology of Myrmeleontoidea also provides an important set of characters that are phylogenetically informative 24 . So far, no definite larva of Babinskaiidae has been reported. Badano et al. 25 presented an interesting study focusing on the higher phylogeny of Myrmeleontiformia by using larval characters from both extant and fossil taxa. Notably, most fossil taxa sampled for the phylogenetic analysis in Badano et al. 25 are from Kachin amber. Thus, this work is highly relevant to the present study and may provide additional evidence to infer the phylogenetic position of Babinskaiidae. Babinskaiidae is the most diverse and abundant group of Myrmeleontoidea from Kachin amber 1 . In the phylogeny recovered in Badano et al. 25 , the diverse Kachin amber myrmeleontoids without definite familial affiliations (e.g., Electrocaptivus xui Badano, Engel and Wang in Badano et al. 25 , Burmitus tubulifer Badano, Engel and Wang in Badano et al. 25 , Adelpholeon lithophorus Badano and Engel in Badano et al., 2018 25 , etc.) are assigned to be the stem-group Ascalaphidae + Myrmeleontidae, while there is no stem-group taxon or "side branch" found for Nymphidae as mentioned in Makarkin et al. 7,21 . Taphonomically, the diverse Kachin amber myrmeleontoid larvae that do not belong to any extant family may be associated with the extinct family Babinskaiidae, which is also diverse in species from the same deposit. Following this assumption, in the phylogenetic tree of Badano et al. 24 , Babinskaiidae is nested within the clade including Nemopteridae, Ascalaphidae, and Myrmeleontidae, which is partially concordant to the present result, although the identical position and monophyly of Babinskaiidae is not recovered as in our result. Of course, there are other possibilities on the identity of these larvae, such as those of the extinct Araripeneurinae that are also relatively basal to the extant antlions. Future in-depth study on the taxonomy of the Kachin amber myrmeleontoid larvae and their associations with the adult forms is important to resolve the deep phylogeny of Myrmeleontoidea. www.nature.com/scientificreports/ currently comprising nine genera and 10 species, display diverse morphological characters (Table S1). Pseudobabinskaia martinsnetoi (Lu et al.) 6 is known to date as the smallest species of the family in body-size (forewing length ~ 9.0 mm), while C. xiai sp. nov. and Gigantobabinskaia godunkoi Makarkin and Staniczek 10 are among the largest babinskaiids (forewing length over 22.0 mm) 6,10 . Specialized wing shape is also remarkable in some Kachin amber babinskaiids. For example, C. xiai sp. nov. and S. punctata sp. nov. possess distinctly narrow and elongate wings; E. burmana, G. godunkoi, and X. lepidotricha sp. nov. have modified hind wings that are distally falcate; and more peculiarly Burmobabinskaia tenuis Lu et al. 6 has strongly narrowed hind wing similar to that of Nemopteridae 6,10 .

Diversification of
The morphological diversity of legs are highlighted by the differently modified tarsi and arolium. The tarsomeres are usually dilated in many babinskaiids, being indicative of an arboreal life mode, but the degree of development greatly varies among species 10 . In G. godunkoi all five metatarsomeres are strongly dilated with long hairs 10 . A similar trait is also present in E. burmana, but being slightly less developed 6 . In the other Burmese amber babinskaiids, such tarsal dilation is not as distinct as the former two species. However, in C. xiai sp. nov. the tarsomeres of all legs are densely setose and bear several long spinous, distally directed setae on the ventral surface. More interestingly, this species has bifid arolium in all legs, suggesting that this character state previously considered to be an autapomorphy of Nymphidae 26 is convergently derived in other myrmeleontoid species. Additionally, in C. xiai sp. nov. there are some regularly spaced spinous setae on femora and tibiae, which seemingly function for predation or defense.
The scaly setae on the basal section of forewing costa in X. lepidotrica sp. nov. is another striking trait (Figs. 7,  8). This is not only the first report of scales in Babinskaiidae but also in all fossil and extant myrmeleontoids. Such scales especially resemble those in some beaded lacewings (Neuroptera: Berothidae). So far, function of the scales in Berothidae is completely unknown, however, in this family these scales are only found in females [27][28][29] . Thus, the scales in female beaded lacewings may be a kind of organs related to chemical communication for attracting or searching mates. Notably, the scales in babinskaiids herein reported are also present in female. Although the male of X. lepidotrica sp. nov. has not been found, if the scales are absent in males of this species, similar function of this trait involved in courtship as hypothesized in Berothidae may be inferred. Nevertheless, convincing evidence awaits discovery. Scales are also present on the wing eyespot of some derived subfamilies of Kalligrammatidae, which is thought to have certain optical function 30 . It is evident that they may be not similar in form and function to the presently reported scales in Babinskaiidae. In the nemopterid subfamily Crocinae, some extant species have a specialized structure called "bulla", which is a group of silky hairs, sometimes ended as a small knob, on the fore-or hind wings 31 . This trait is confined to males and thought to be a kind of scent organ 31 . Similarly, the pilula axillaris or Eltringham organ is a club-like projection laterally with a tuft of setae from the hind wing margin, and it is also confined to male antlions. However, this structure is proposed to be a dispersing organ that could spread but not produce the sexual pheromones 32 . In some fossil species of the extinct chrysopoid family Mesochrysopidae there is a bunch of elongated hairs on the forewing 33 . Possibly, this trait also has a similar function as the scales or specialized hairs or organ in aforementioned myrmeleontoids, although whether it is present in only male or female or both sexes is unknown.
Finally, it is noteworthy to mention some previously unknown characters of the female genitalia in Babinskaiidae. In C. xiai sp. nov. the valvate female gonocoxites 9 bear some short dentoid processes along the ventral margin. In many antlions there are a number of stiff setae called digging setae on some female genital sclerites, such as gonocoxites 8, gonocoxites 9, and ectoprocts 34,35 . This is an adaptive trait for laying eggs on sand-like substrate 34,35 , which is probably associated with the fossorial life-style of some antlion larvae. In X. lepidotrica sp. nov. there is an additional lobe on anteroventral portion of the female gonocoxite 9, and this modification is similar to the hypocaudae in some berothid species 28 . In total, five types of female genitalia have thus far been described in Babinskaiidae (see Lu et al. 6 ; Hu et al. 8 ), suggesting disparate copulation or oviposition behaviors of these archaic myrmeleontoid species.

Conclusion
The palaeofauna of Babinskaiidae from the mid-Cretaceous Kachin amber of Myanmar is represented by diverse genera and species with disparate morphological modifications. Our findings provide significant new morphological evidence for understanding the phylogenetic position and diversity of Babinskaiidae. In terms of the phylogenetic position of Babinskaiidae herein recovered, this Cretaceous family probably represents a transitional lineage between Nymphidae and the advanced myrmeleontoids. The Cretaceous radiation and recent decline of Babinskaiidae remains mysterious, awaiting discovery of palaeodiversity and phylogenetic study combining adult and larval characters.

Material and methods
Taxonomy. The 36 : Fig. 1). The age of this deposit is dated to be ~ 99 million years (the earliest Cenomanian) by U-Pb dating of zircons from the volcaniclastic matrix of the amber 37 .
The type specimens are deposited in the Nanjing Institute of Geology and Palaeontology (NIGP), Chinese Academy of Sciences, Nanjing; the Lingpoge Amber Museum (LAM), Shanghai; and the Century Amber Museum (CAM), Shenzhen.
All taxonomic acts established in this paper have been registered in ZooBank, together with the electronic publication: urn:lsid:zoobank.org:pub:605A4DCC-9ADE-473C-93BD-31878E03FA07. Phylogenetic analysis. Aiming to reveal the phylogenetic position of Babinskaiidae within Myrmeleontoidea, a phylogenetic analysis was performed based on samples from major fossil and extant lineages of Myrmeleontoidea. For the ingroup taxa in the present analysis, we sampled most known genera of Babinskaiidae except Baisonelia (only hind wings preserved) and Pseudoneliana (distal part of both wings not preserved). For Nymphidae, the stem-group of this family (Liminympha Ren and Engel 41 ) as well as the extant representatives of Nymphinae (Nesydrion Gerstaecker 42 and Nymphes Leach 43 ) and Miodactylinae (Osmylops Banks 44 and Norfolius Navás 45 ) were selected. Two nymphid-like genera from the Lower Cretaceous of Brazil, i.e., Araripenymphes and Cratosmylus, which were considered to represent a separate family Cratosmylidae by Makarkin et al. 7 , were selected. Three genera of the extinct family Palaeoleontidae, i.e., Baisopardus Ponomarenko 4 , Paraneurastenyx Martins-Neto 46 , and Parapalaeoleon Menon and Makarkin 47 , were selected because the species of these genera have better preserved morphological characters than the other species of this poorly known myrmeleontoid fossil family. The other sampled ingroup taxa are the same to those in Lu et al. 22 . Notably, the genera Stilbopteryx and Palpares are assigned to the subfamily Ascalaphinae in Machado et al. 48 , representing ascalaphids in the present sampling. We selected Gumilla Navás 49 (Osmylidae), Ithone Newman 50 (Ithonidae) and Balmes Navás 51 (Psychopsidae) as the outgroup taxa.
Morphological characters used in the phylogenetic analysis are mainly from Martins-Neto 13,14 and Lu et al. 22 with some modification, comprising a total of 54 adult characters (see Note S1). Unknown characters were coded as "?", while inapplicable characters were coded as "-". The data matrix is also given in Table S2. All characters were treated as unordered and with equal weight. We analyzed the dataset using TNT v1.5 52 with an initial Traditional search (Starting trees: 100 repls, TBR, trees to save per replication: 10). Bremer support values and Bootstrap values were calculated with the function implemented in TNT (Setting for Bremer support values calculation: TBR from existing trees, retain trees suboptimal by 10 steps; setting for Bootstrap values calculation: traditional search, number of replicates: 1000). Character states were mapped on the strict consensus tree (MPT) using WinClada ver. 1.00.08 53 , showing only unambiguous changes.