Doratomantispinae Lu et al., 2020

Subfamily Doratomantispinae Lu et al., 2020

Revised diagnosis Vertex posteriorly with two lateral tubercles. Pronotum tubular, with paratergal lobes ventrally fused; maculae absent; postfurcasternum very small. Profemur distinctly shorter than protibia plus protarsi, not compressed along ISs row (without posteroventral carina), distinctly broadened near base, ventrally with two ISs rows: ISs stout, saber-shaped, rarely spine-shaped; anteroventral row consisting of a major process with two, four, or five basal branches each distally bearing a conical Stitz organ, and several short processes along distal 1/3–3/4 length of profemur; posteroventral row with 3–18 processes, commencing slightly distal to level of major process. Protibia ventrally with a row of flattened (Type j6, j9) or angularly curved (near Type k4) prostrate setae on well-developed and thickened keel; protarsus 5-segmented, tarsomere 1 not prolonged and not distally spinously produced, tarsomere 1–4 ventrally bearing conical setae (Type k3); two simple pretarsal claws; arolium small, subtriangular, acutely tapering. Wing trichosors present almost along entire wing margin, mostly with just one between adjacent veinlets; ScP distally abruptly bending towards and fused with RA, pterostigma present between C and ScP + RA, one or two series of gradates (inner gradates incomplete) present; forewing CuP not approximating A1 proximally; hind wing 1rp-m upright, CuP reduced with stem weak or absent. Male gonocoxites 9 short or long, slender, or broad, distally simple or bifurcated; gonapophyses 10 long, slender, or broad, distally acute; gonostyli 10 (pseudopenis) whip-shaped, short to very long, basally enlarged and thickened. Female tergum 9 not fused with ectoprocts; pseudohypocaudae absent; gonocoxites 9 short, not elongate into a long ovipositor.

Key to genera of Doratomantispinae

1. Major process with two basal branches (anterior branch absent). Protibia with a row of angularly curved prostrate setae (near Type k4). Male gonocoxites 9 short and broad; gonapophyses 10 blade-like. Female sternum 7 posterolaterally not produced; ectoprocts ventrally produced……….………....…… Doratomantispa Poinar in Poinar & Buckley

– Major process with four or five basal branches (anterior branch present). Protibia with a row of flattened prostrate setae (Type j6, j9). Male gonocoxites 9 long and slender; gonapophyses 10 rod-like. Female sternum 7 posterolaterally produced; ectoprocts ventrally not produced….. Paradoxomantispa Lu et al.

The adult morphology of Doratomantispinae

Body size Doratomantispinae are small to median sized mantispids with a body length (BL) ranging from 4.38 to 15.25 mm and forewing length (LFW) ranging from 3.86 to 12.06 mm. Doratomantispa (LFW: 6.88–12.06 mm, BL: 9.00– 15.25 mm; the largest species: D. ares with LFW 12.06 and BL 15.25 mm, the smallest species: D. zhuozhengmingi sp. nov. with LFW 6.88 and BL 8.51 mm) is generally larger in body size than Paradoxomantispa (LFW: 3.86–5.64 mm, BL: 4.38–5.8 mm; the largest species: P. jiaxiaoae with LFW 5.64 and BL 6.29 mm, the smallest species: P. mahaiyingae sp. nov. with LFW 3.86 and BL 4.38 mm). The adult body size in extant Mantispidae (including Symphrasinae ) shows remarkable inter- and intra-specific variations, being less useful for species determination and possibly associated with the amount of food ingested by the larvae ( Ardila-Camacho et al., 2021; Hoffman, 1992; Lambkin, 1986a, b; Redborg, 1998; Snyman et al., 2020). Although the variation of body size of the male specimens of P. mahaiyingae sp. nov. is not distinct, we at present could not determine whether the body size is useful for the species identification in Doratomantispinae , as many species of this subfamily are only described based on a single specimen.

Head The vertex of Doratomantispinae bears two large lateral tubercles posteriad the distinctly broadly raised supra-antennal area, medially with a distinct coronal suture extending to the supra-antennal area or restricted on its narrow posterior portion ( Figs. 5 View Fig d-e, 8c, 12e-f, 15b-c, 17d-e, 22a-b). This feature is also found in Paraberothinae and some berothids (e.g., Berotha , Nosybus , Podallea ), but tubercles in these taxa are generally located at or slightly anteriad the posterior end of the broadly raised supra-antennal area and their coronal suture seems not as developed as in Doratomantispinae . Notably, some authors consider this broadly raised supra-antennal area as one median tubercle (McLeod & Adams, 1967). In Mantispoidea, the vertexal tubercles are usually developed in Berothidae and Rhachiberothidae , being modified as two (see above) or three tubercles (excluding the case of the broadly raised supra-antennal area regarded as the median tubercle) (e.g., in Lekrugeria , Rhachiberotha ) or being fused into a large tubercle, such as in Mucroberotha . However, in Mantispidae , only Doratomantispinae and Psilomantispa have the paired, developed lateral vertexal tubercles, and Calomantispa venusta is the only extant species having two small and less developed anterolateral tubercles on the vertex. In the other mantispids, the supra-antenal area raises above the toruli as two protuberances (e.g., Plega ) or one medial tubercle-like hump (e.g., Gerstaeckerella and some mantispines). The vertexal region of all dipteromantispids lack tubercles, with supra-antennal area not raised but as smooth as in some symphrasines (e.g., Trichoscelia and some species of Anchieta ) ( Ardila-Camacho et al., 2021; Li & Liu, 2020; Li et al., 2020b; Liu et al., 2016).

Prothorax Being similar to Drepanicinae , Calomantispinae , and Mantispinae , the pronotum in Doratomantipinae is tubular, with the paratergal lobes fused ventrally and the distinctly expanded anterior end where the forelegs insert ( Figs. 1 View Fig a-b, 5b-c, 7b-c, 12b, 13a, 15b-c, 17b, 20b, 23c); the basisternum is anteriorly connected to the episternum through the precoxal bridge (although only visible in D. yumeiyingae sp. nov. ( Fig. 13b View Fig )); the postfurcasterna is present as two narrow posterolateral sclerites ( Ardila-Camacho et al., 2021) ( Fig. 17c View Fig ). All doratomantispines lack the pronotal maculae, which is present as a pair of shagreened or smooth and glossy spots or small humps with many squamous or reticular sensilla, located at the base of pronotal anterior expanded portion in the three mantispid subfamilies aforementioned ( Ardila-Camacho et al., 2021; Lambkin, 1986a; Li et al., 2020a; fig. 4I-L; Poivre, 1978; fig. 2E). Setae on the doratomantispanie pronotum are pedicellate, long, and thick, resembling those in some symphrasines (e.g., Plega ), some drepanicines (e.g., Allomantispa ) and calomantispines, and they are sometimes lost only with the pedicellate bases left in amber ( Figs. 8a View Fig , 13a View Fig , 17 View Fig b-c, 20b, 23c; Ardila-Camacho et al., 2021; Lambkin, 1986a, b).

Raptorial foreleg As Rhachiberothidae , Symphracinae, and Calomantispinae , the procoxal surface of Doratomantispinae is smooth, lacking the posterodistal impressed (present in Drepanicinae ) or flattened (present in Drepanicinae and Mantispinae ) area, or with a proximal annular groove (present in Mantispinae ) ( Ardila-Camacho et al., 2021; Lambkin, 1986a, b; Li et al., 2020a). The protrochanter of Doratomantispinae is similar to that of Drepanicinae and Calomantispinae , being subconical with the “trochanterfemur complex” association less developed than in Mantispinae ( Ardila-Camacho et al., 2021; Büsse et al., 2021).

The profemur of Doratomantispinae ( Figs. 2 View Fig a-b, e-f, 5h-i, 7f, 8d-e, 11a, c-d, 13c-d, 15d, g, 20c-d, 22c-e, j, 23e, 24a) varies from robust to slender, being distinctly broadened near base and narrowed towards the proximal and distal ends respectively (i.e., with a distinct ventral angle near base). This feature is very distinctive in Mantispidae (generally broadened near middle) but similar to some dipteromantispids (e.g., Burmodipteromantispa ) and some rhachiberothids (e.g., Acanthoberotha , Creagroparaberotha , Mucroberotha , Raptorapax , Rhachiella , Uranoberotha ). The profemoral integumentary specializations of Doratomantispinae are distinctly reclined forwards with the large angle of inclination as Drepanicinae , Calomantispinae , and Mantispinae , and they possess some remarkable characters in the raptorial Mantispoidea as follows:

1. Integumentary specializations (ISs) developed, stout, and saber-shaped ( Ardila-Camacho et al., 2021: fig. 29, Type c1-6, 8–9; few spine-shaped ISs only found in D. yumeiyingae sp. nov.). By contrast, the ISs have the following different modifications: long and slender, mainly stinger-shaped and thorn-shaped in Paraberothinae (sometimes the end of profemur with short saber-shaped or tubercle-shaped ISs), Rhachiberotha and Rhachiella ; slender, mainly saber-shaped with several thorn-shape ISs near the profemoral base in Paradoxoberotha ; mostly very short, stinger-shaped, thorn-shaped, and tubercle-shaped, with several long and thorn-shaped ISs in Mucroberotha and Whalfera ; all tubercle-shaped in Hoelzeliella and Dipteromantispidae ; very short and possibly all tubercle-shaped in Mesomantispinae ; very short, mainly tubercle-shaped in Symphrasinae (Some stinger-shaped and spine-shaped ISs also occur in certain genera. Besides, there are another two rows of thickened setae each with a glabrous asymmetrical base in the subfamily); long, spine-shaped alternating with some short spine-shaped or stinger-shaped ISs in Drepanicinae , Calomantispinae , and Mantispinae ; many stinger-shaped mixed with several long saber-shaped ISs, and distal ones short saber-shaped in Aragomantispa ; all disappeared in Psilomantispa . All the ISs are also saber-shaped in Acanthomantispa , Dicranomantispa , and Pectispina , but much more reduced and fewer than those in Doratomantispinae (see the terms and types of ISs in Ardila-Camacho et al. (2021: fig. 29, table. 1)).

2. The presence of basally branched major process. The branched specialization in Doratomantispinae , Acanthomantispa , Dicranomantispa , and Pectispina is here considered to be homologous to the major process in extant Drepanicinae , Calomantispinae , and Mantispinae , based on this structure long, situated most basally at the anteroventral row and the mantispid affinity of these taxa. The branched major process differs from the simple one in Mantispidae as each branch distally bears a conical Stitz organ rather than plug-shaped, namely each branch saber-shaped rather than spine-shaped. The major process in Doratomantispinae can be distinguished from that in Acanthomantispa , Dicranomantispa and Pectispina by the primary branching point at base (primary branching point near the proximal 1/6–1/5 length of major process in Acanthomantispa and Pectispina , at the middle of major process in Dicranomantispa ). There is also a stout and longest process similar to the major process near the base of anteroventral row in some rhachiberothids, Anchieta and Plega , e.g., the stout and longest thorn-shaped process in Astioberotha , Micromantispa , Stygioberotha , and Paradoxoberotha , and the stout and longest spine-shaped process in Anchieta and Plega , which may be convergent to the major process. The length of the major process in Doratomantispinae (excluding P. mahaiyingae sp. nov. (only 1/3 length of protibia)) is very long, mostly 1/2 to nearly the same length of protibia. The similar state is also found in Acanthomantispa and Pectispina , while it is less elongated in the other raptorial Mantispoidea (see the terms and types of ISs in Ardila-Camacho et al. (2021, fig. 29, table. 1)).

3. The anteroventral row in Doratomantispinae is reduced to the basal branched major process and several short processes generally gradually shortened towards the profemoral end (only not in D. gaoyuhei sp. nov.), being arranged sparsely along the profemoral distal 1/3–1/ 2 in Doratomantispa or distal 1/2–3/ 4 in Paradoxomantispa . By contrast, in the mantispoids with major process or its convergent process, the short processes of anteroventral row are always not gradually shortened and arranged more densely and/or more apically in some rhachiberothids and symphrasines (e.g., species in Astioberotha , Mucroberotha , Paradoxoberotha , Rhachiberotha ; Anchieta , Plega ), and are almost completely lost in Drepanicinae , Calomantispinae , and Mantispinae , except some genera, such as Gerstaeckerella and Nolima , possessing fewer short processes at the apex of profemur ( Ardila-Camacho et al., 2021, figs. 26d, 27a, b; Aspöck & Aspöck, 1997, fig. 1; Aspöck & Mansell, 1994, fig. 11; Nakamine et al., 2020, 2021; Penny, 1982, figs. 11–12).

4. Compared with other mantispoids except some drepanicines and its related genera, the processes of posteroventral row in Doratomantispinae are arranged more sparsely, and completely or nearly gradually shortened towards the profemoral end, not alternating with short or minute processes, except P. jiaxiaoae and P. mahaiyingae sp. nov.. This row of processes is usually well developed in the raptorial Mantispoidea, but extremely reduced or even absent in a few Burmese amber mantispids (e.g., Acanthomantispa , Pectispina , Dicranomantispa , and Psilomantispa ) ( Lu et al., 2020; Shi et al., 2020).

Unlike the smoothly to strongly curved protibia of other raptorial mantispoids (excluding some paraberothines), that of Doratomantispinae ( Figs. 2 View Fig a-b, e-f, 5h-i, 7f, 8d-e, 11a, c-d, 13c-d, 15d, g, 20c, d, 22c-e, j, 23e, 24a) is as straight as that of an ambulatory foreleg, but still distinctly curved at the joint with the profemur. The protibial distal margin of this subfamily is not acutely produced, without tibial spur, and the ventral keel is well developed and thickened. The protibial prostrate setae of Paradoxomantispa are flattened and arranged closely, being identical to those in Symphrasinae , extant Drepanicinae , Calomantispinae , and Mantispinae ( Figs. 22g, j View Fig , 23h View ◂ , 24a View ◂ ), while those of Doratomantispa are distally angularly curved and arranged sparsely, which is unique in Manrtispoidea and may be an apomorphy of the genus ( Figs. 1h View Fig , 5i View Fig , 8f View Fig , 11a View Fig , c-d, 13f, 15e, 17g, 20f). Notably, the protibial prostrate setae of Acanthomantispa is also arranged sparsely with their base upright and apex feebly curved, being similar to those of D. zhuozhengmingi sp. nov., D. gaoyuhei sp. nov. and D. pubescens, ( Lu et al., 2020) . The protarsus of Doratomantispinae appears to be generalized as in most raptorial Mantispoidea except Symphrasinae and some rhachiberothids. The protarsal ventral conical setae and the small subtriangular arolium are very distinctive in the raptorial Mantispoidea, and only found in Doratomantispinae , Acanthomantispa , Dicranomantispa , Pectispina , and Psilomantispa ( Lu et al., 2020; Shi et al., 2020).

Wings The wings of Doratomantispinae possess some characters common or typical in Berothidae and basal raptorial mantispoid groups, but weakly developed or absent in the derived mantispid clade including Drepanicinae , Calomantispinae , and Mantispinae . First, in Doratomantispinae , the trichosors are present along the entire wing margin except its basal portion, and there is only one trichosor between adjacent veinlets although sometimes two to three trichosors sporadically are present on the anterior wing margin ( Figs. 3a View Fig , 18 View Fig a-b, 21a). Similar state is also present in Berothidae , Rhachiberothidae , Mesomantispinae , and some fossil genera of Symphrasinae ( Archaeosymphrasis , Habrosymphrasis , Parasymphrasites ).

Second, the forewing costal space of most doratomantispine species bears some or many continuously arranged branched costal crossveins, which is similar to some berothids (e.g., extant berothids, Elektroberotha , Maculaberotha ), Rhachiberothidae , some fossil symphrasines (e.g., Archaeosymphrasis , Parasymphrasites ), and Mesomantispinae , while the forewing costal crossveins are generally all simple in Drepanicinae , Calomantispinae , and Mantispinae , except some species of Drepanicus and Gerstaeckerella .

Third, the ScP is distally fused with RA as in some fossil berothids (e.g., Ansoberotha , Cantabroberotha , Cornoberotha , Dolichoberotha , Maculaberotha , Plesiorobius , Sibelliberotha ), Paraberothinae , Paradoxoberotha , some mesomantispines (e.g., Ovalofemora ), and Symphrasinae , while in Drepanicinae , Calomantispinae , and Mantispinae , the ScP separates from RA by a narrow or slightly broad hyaline stripe with one or more crossveins.

Fourth, the hind wing CuA of Doratomantispinae is long and pectinately branched into multiple veins (7–12 main branches, all simple or largely bifurcated), terminating at or distad the midpoint of wing posterior margin, which is similar to Berothidae , Rhachiberothidae , Symphrasinae , some fossil drepanicines ( Acanthomantispa , Dicranomantispa ), and their related genus Pectispina . By contrast, generally in the derived mantispid clade, the hind wing CuA is short, pectinately branched, or forked, terminating distinctly proximad the midpoint of wing posterior margin, sometimes rather reduced in Calomantispinae and Mantispinae .

Notably, it is charactistic for Mantispoidea that the hind wing 1rp-m of almost all doratomantispines occurs distal to the primary branching point of RP and at or proximal to that of M (slightly distal to that of M only in D. burmanica ). By contrast, in remaining mantispoids, if this hind wing crossvein present, it is commonly proximal to the primary branching point both of RP and M in extant berothids, some fossil berothids ( Cantabroberotha , Sibelliberotha ), Rhachiberothidae , Symphrasinae , Gerstaeckerella , Theristria , and Nolima ; proximal to the primary branching point of RP and slightly to distinctly distal to that of M in Nyrma , some fossil berothids ( Ansoberotha , Cornoberotha , Dolichoberotha , Elektroberotha , Systenoberotha ), Rhachibermissa ( Paraberothinae ), Allomantispa , Calomantispa , and Mantispinae : distal to the primary branching point both of RP and M in Acanthomantispa , Dicranomantispa , Ditaxis , and Drepanicus .

The hind wing CuP of Doratomantispinae still retains the stem parallel to Cu, being vestigial and slightly extending proximad as in Drepanicus ( Figs. 6c View Fig , 14c View Fig , 16 View Fig c-d, 18c, 24f, h), or distinctly diverging from Cu near wing base as in some fossil berothids, Rhachiberothidae and Symphrasinae ( Fig. 3c View Fig ; Ardila-Camacho et al., 2021; Shi et al., 2019b; Fig. 3B View Fig ). The CuP in Doratomantispinae is distally separated from CuA by a short crossvein or directly coalesced with it at a point, rather than largely fused with it as in Rhachiberothinae , because the CuP distally possesses the definitive branch and runs towards CuA, forming a loop, which is very similar to the case in Drepanicus but different from that in Rhachiberothinae ( Ardila-Camacho et al., 2021; Breitkreuz et al., 2017). Additionally, the hind wing CuP of Doratomantispinae is probably separated distally from A1. In contrast to the taxa with hind wing CuP proximally parallel to Cu and distally fused with A1 ( Rhachiberothinae and Drepanicus ), the vein connecting CuP with A1 at their fusion point is located on the branch of A1, which belongs to a part of CuP or anterior A1 branch. However, in Doratomantispinae , the similar vein is usually located at the stem of A1 as in some extinct symphrasines ( Archaeosymphrasis , Habrosymphrasis ) ( Figs. 3c View Fig , 14c View Fig , 16 View Fig c-d, 18c; Lu et al., 2020, figs. S4I; Shi et al., 2019a, fig. 3B), indicating that it might be a real crossvein linking CuP with A1 ( Ardila-Camacho et al., 2021; Shi et al., 2019b). The hind wing CuP as in Doratomantispinae also occurs in some extinct berothids ( Ansoberotha , Cornoberotha , and Dolichoberotha ) and drepanicines ( Acanthomantispa , Dicranomantispa , and Pectispina ) ( Lu et al., 2020; Shi et al., 2020; Yang et al., 2020).

Male terminalia ( Figs. 4 View Fig , 6 View Fig d-e, 16e-f, 19, 21d-e, 25; Lu et al., 2020, S4D-G; Shi et al., 2020, 5C-D). The tergum 9 is half-ring shaped, separated from ectoprocts, very narrow to broad, dorsally generally narrower than lateral and ventral portions, being similar to other raptorial mantispoids and some berothids ( Cornoberotha , Cyrenoberotha , Dolichoberotha , Manselliberotha ) ( Ardila-Camacho et al., 2021; Aspöck & Aspöck, 1988b; McLeod & Adams, 1967; Yang et al., 2020). The sternum 9 is not fused with sternum 8, with the shape and the position relative to ectoprocts similar to that in some species of Drepanicinae and Mantispinae . However, in D. arcimaculata sp. nov., the sternum 9 is similar to that of Calomantispinae by its scoop-shaped, remarkably enlarged, and elongated condition ( Ardila-Camacho et al., 2021; Lambkin, 1986b; Reynoso-Velascol & Contreras-Ramos, 2019). The ectoprocts are completely paired, close to each other, ovoid with variable length, with distinct callus cerci visible in some well-preserved specimens, which is similar to that in Calomantispa venusta , Drepanicus , and some species of Gerstaeckerella and Theristria ( Ardila-Camacho et al., 2021; Lambkin, 1986a, b; Liu et al., 2015; Poivre, 1978).

Besides, the male genital sclerites in Doratomantispinae appear more similar to the typical symphrasine configuration in the distinctly tripartite gonocoxites + gonapophyses + gonostyli 10 complex (at least the gonapophyses 10 conspicuously separated from the gonostyli 10), the whip-shaped gonostyli 10 with a thickened and broadened base, and the long and developed gonapophyses 10. It is noteworthy that the gonostyli 10 in Doratomantispinae as in Mucroberotha and Dipteromantispidae has its base curved dorsad and the whip-shaped portion largely extending out of the genital chamber, while in Symphrasinae , this structure is straight or has its base curved ventrad and the whip-shape portion mainly occurs towards the anterior abdominal part though its apex protrudes out from the abdomen ( Ardila-Camacho et al., 2021; Liu et al., 2017). Additionally, in Rhachiella , the gonostyli 10 (fused with gonocoxites 10 and gonapophyses 10 in this genus) has its base curved ventrad as in Symphrasinae , but the whip-shaped portion is curved dorsad near its basal 1/3, being U-shaped and present at the terminal part of the genital chamber ( Aspöck et al., 2020, figs. 2, 5–6). The genital sclerites in Paradoxomantispa share another two characters with Symphrasinae : the paired, slender, rod-like gonocoxites 9 each with an acute apex (present in some species of Anchieta and Plega spinosa Ardila-Camacho et al., 2019 ) and the paired, slender rod-like gonapophyses 10 ( Ardila-Camacho et al., 2021). In Doratomantispa , the gonocoxites 9 are similar to Drepanicinae in the sclerites very short, stout and broadened, distally obtuse or present as one or two acute hook-like processes, and the gonapophyses 10 are long blade-like, anteriorly weakly sclerotized, tapering posteriorly into a strongly sclerotized point. The gonocoxites 11 in Doratomantispinae posteriorly bears a median lobe as in other mantispids but its entire shape still remains unclear, being only known as arch-like anterodorsally with two lateral slender branches in D. longa (similar branches also found in some berothids, e.g., Asadeteva , Berotha ). The median lobe of gonocoxites 11 in the subfamily shows great variations, being short, broad, posteriorly obtuse, or acutely produced, sometimes dorsally with four spines in Doratomantispa , while long and medially deeply concaved dorsad in Paradoxomantispa .

Female terminalia The configuration and morphology of visible portions of female terminalia in the subfamily present similarity to some drepanicines, Calomantispinae and Mantispinae . It is noteworthy that the sternum 7 in Paradoxomantispa posteriorly bears two distinct lateral projections very similar to those of Dipteromantispidae ( Li & Liu, 2020; figs. 1E-F; Liu et al., 2016; figs. 9–10; Lu et al., 2020, figs. S6E-F), and the gonocoxites 9 or ectoprocts are produced ventrally as in some species of Theristra Lambkin (1986a).

Consideration on the possible phylogenetic position of Doratomantispinae

In the phylogenetic analysis of Lu et al. (2020), two autapomorphies of Doratomantispinae were discovered, i.e., (1) a profemoral processes gradually shortened distad, and (2) a subtriangular, profemoral arolium with an acutely tapering apex. After a comprehensive comparison based on new fossils, we found some exceptions in Doratomantispinae with profemoral anteroventral or posteroventral processes arranged irregularly, such as in D. gaoyuhei sp. nov., P. jiaxiaoae , and P. mahaiyingae sp. nov.. Character (2) is also shared by some extinct drepanicines and their related genera (e.g., Acanthomantispa , Dicranomantispa , and Pectispina ) ( Lu et al., 2020, figs. S12E-L; Shi et al., 2020, 3D-E). Moreover, we found five more possible synapomorphies of this subfamily, including (1) the vertex with two large lateral tubercles posteriad the broadly raised supra-antennal area, (2) the profemoral ISs stout and almost always saber-shaped, (3) the anteroventral row sparse, arranged along the distal 1/3–3/4 length of the profemur, (4) the major process branched basally, and (5) the hind wing 1rp-m distal to the primary branching point of RP and at or proximal to that of M.

The familial and subfamilial relationships within Mantispoidea are controversial as indicated in recent phylogenetic studies ( Ardila-Camacho et al., 2021; Liu et al., 2015; Lu et al., 2020; Shi et al., 2019b, 2020; Winterton et al., 2018). In particular, based on the basic phylogenomic data, Winterton et al. (2018) recovered Mantispidae as a paraphyletic group due to the sister relationship between Symphrasinae and Rhachiberothidae as well as the sister relationship between the derived mantispid clade and Berothidae . Ardila-Camacho et al. (2021) further supported the rhachiberothid affinity of Symphrasinae and transferred this subfamily to Rhachiberothidae based on a morphology-based phylogeny, but they recovered Rhachiberothidae + Mantispidae as a monophyletic group—raptorial Mantispoidea as the sister to Berothidae . However, these two studies lacked sampling of sufficient berothid groups and a broad array of fossil taxa. The disputed rhachiberothid affinity of Symphrasinae causes confusion on the phylogenetic position of Doratomantispinae , which was previously recovered as the transitional linage between Symphrasinae and the derived mantispid clade ( Liu et al., 2015; Lu et al., 2020) or as the sister to the former ( Shi et al., 2020).

The present study provides additional evidence concerning the phylogenetic placement of the Doratomantispinae . First, this subfamily certainly belongs to the raptorial Mantispoidea by having the synapomorphies of this group: (1) the presence of a distinct, well-developed postfurcasternum, (2) raptorial foreleg, with incrassate femur, (3) the presence of Stitz organs, (4) two rows of profemoral integumentary specializations, and (5) the keeled protibia ventrally with specialized setae ( Ardila-Camacho et al., 2021). Second, we support that this subfamily has closer affinity to the derived mantispid clade, assigned to Mantispidae as recovered in Lu et al. (2020), based on the following putative apomorphic states: (1) ocular plane anteriorly curved, (2) pronotum tubular with paratergal lobes ventrally fused and expanded area of procoxal insertion, (3) foreleg inserted at the anterior apex of prothorax, (4) the presence of precoxal bridge, and (5) the presence of a narrow postfurcastenum ( Ardila-Camacho et al., 2021). Doratomantispinae possesses some characters shared with Symphrasinae and other basal raptorial mantispoids, i.e., (1) the absence of maculae, (2) the well-developed trichosors almost along the entire wing margin, (3) the frequent development of branched costal crossveins, (4) ScP distally fused with RA, (5) hind wing CuA terminating at or distad to the midpoint of the wings posterior margin, and (6) gonostyli 10 basally broadened and thickened. All of these features, however, appear to be plesiomorphic. Notably, there are some characters only present in Symphrasinae and Doratomantispinae , i.e., the long and developed gonapophyses 10, the presence of distinctly tripartite gonocoxites + gonapophyses + gonostyli 10 complex, and the paired long, slender, rod-like gonocoxites 9 and gonapophyses 10 (only known in Paradoxomantispa in Doratomantispinae ). Considering the uncertain familial placement of Symphrasinae and the lack of knowledge of the male genitalia in many fossil raptorial mantispoids, it is still difficult to determine whether those common characters evolved independently in these two subfamilies or represent a plesiomorphic state in the raptorial Mantispoidea.