﻿Six new species of Margattea Shelford, 1911 (Blaberoidea, Pseudophyllodromiidae, Neoblattellini) from China

﻿Abstract Six Margattea species are established and described: three are cryptic species, namely, M.parabisignata Li & Che, sp. nov., M.semicircularis Li & Che, sp. nov., and M.forcipata Li & Che, sp. nov. They are distinguished from known species M.bisignata, M.spinifera, and M.paratransversa by their male genitalia with the aid of molecular species delimitation method (ABGD) using COI as the molecular marker. The other three new species are M.pedata Li & Che, sp. nov., M.undulata Li & Che, sp. nov., and M.bisphaerica Li & Che, sp. nov. Morphological and genitalia photographs of these new species of Margattea, as well as a key to the species of Margattea from China, are provided.


Introduction
A total of 63 species of the genus Margattea in Neoblattellini have been recorded in Asia, Africa, and parts of Oceania (Beccaloni 2014), and 26 of them are known from China (He et al. 2021).There are twelve genera in Neoblattellini.Margattea is the only genus that is not distributed in the New World, while all the other eleven genera are only present in the New World.Margattea is morphologically similar to the genus Balta in the yellowish brown body, the subelliptical pronotum (sometimes with spots), and wings that usually exceed the end of the abdomen (He et al. 2021).Despite being a relatively well-diversified genus, only four molecular phylogenetic analyses sampled Margattea species (Evangelista et al. 2021(Evangelista et al. , 2023;;Liu et al. 2023;Wang et al. 2023).Due to the different molecular data and sampling taxa in molecular analyses listed above, the sister group of Margattea was inconsistently recovered, but Balta was always found phylogenetically distant from Margattea.
At present, members of Margattea are identified by the simple, cylindrical, and symmetrical styli, the usually specialized eighth abdominal tergum, the pronotal disc with symmetrical stripes and maculae, and the median phallomere usually with accessory structure (Roth 1989;Wang et al. 2009;Liu and Zhou 2011;He et al. 2021).However, these diagnostic characteristics can be found separately in other genera; moreover, the styli of Margattea are diverse: asymmetrical in Margattea pseudolimbata Wang, Li, Wang & Che, 2014 (Wang et al. 2014), symmetrical and foot-shaped styli or asymmetrical and spherical styli in other samples we examined.Therefore, it is still necessary to further determine the diagnostic characteristics of Margattea.

Morphological study
All type specimens are deposited in College of Plant Protection, Southwest University, Chongqing, China (SWU).Male genital segments were immersed in 10% NaOH solution and incubated with water at 90 °C for 15 minutes to dissolve the fat.All segments were dissected and stored in glycerol for observation, and preserved along with the remainder of the specimen which is stored in ethyl alcohol.All photos were taken by a Leica DFC digital microscope camera attached to a Leica M205A stereomicroscope, then modified with Adobe Photoshop CC 2019.Specimens examined were measured by Vernier Caliper.Morphological terminology mainly follows Roth (2003).The sclerites of male genitalia mainly follows McKittrick (1964).The terminology of veins follows Li et al. (2018)

DNA extraction, amplification, and sequencing
Total DNA was obtained from legs and thoracic muscle using the Hipure Tissue DNA Mini Kit, and the remainder of the specimen was stored in ethyl alcohol.The primers were used to amplify the 658 bp cytochrome c oxidase subunit I (COI) fragment (Table 1).The amplification conditions were: initial denaturation at 98 °C for 2 min, followed by 35 cycles for 10 sec at 98 °C, 10 s at 49-51 °C, and 3 min for 72 °C, with final extension of 3 min at 72 °C, then held at 4 °C.The amplified samples showing bands in agarose gels were sequenced by Beijing Tsingke Biotech Co., Ltd.

Sequence processing and molecular analysis
A total of 105 COI sequences were analyzed, of which 53 sequences are obtained in this study and 52 sequences were downloaded from GenBank (Balta vilis, Sorineuchora nigra, and Mantis religiosa were selected as outgroups) (Table 2).All sequences were aligned by MEGA 7 (Kumar et al. 2016) and adjusted after translation into amino acid sequences.The genetic divergence value was quantified based on Kimura 2-parameter (K2P) (Kimura 1980) by MEGA 7. The Maximum Likelihood (ML) tree was constructed in PhyloSuite v. 1.2.2 (Zhang et al. 2020), using IQ-TREE v. 1.6.8 (Nguyen et al. 2015) with 1,000 ultrafast bootstrap replicates (Hoang et al. 2018).The GTR+G4+F model was selected by ModelFinder 2 (Kalyaanamoorthy et al. 2017) according to the corrected Akaike Information Criterion (AICc).We used a molecular species delimitation method (ABGD: Puillandre et al. 2012) to delimit Margattea species based on COI sequences.ABGD, compared to GMYC and bPTP, provides more conservative estimates, which did not overestimate the number of entities (Puillandre et al. 2012;He et al. 2021).For ABGD, the analysis result was displayed on a web interface (https://bioinfo.mnhn.fr/abi/public/abgd/abgdweb.html).The default parameters were used except for the relative gap width set at 1.0 and using the Jukes-Cantor (JC69) distance.

Morphological species delimitation based on external morphology
Combining the external morphological character, we identified 22 morphospecies of Margattea from a large number of samples collected, including three new species, M. pedata Li & Che,sp. nov.,M. undulata Li & Che,sp. nov.,and M. bisphaerica Li & Che,sp. nov. (Fig. 1A).

Molecular phylogeny and species delimitation based on COI
In this study, we acquired 105 COI sequences of Margattea representing 22 morphospecies of Margattea.The ML phylogenetic tree showed that samples (including males, females, and nymphs) of the same morphospecies form monophyletic groups, although most of the nodes did not have high bootstrap values (Fig. 1).24 molecular operational taxonomic units (MOTUs) were delimited by ABGD (Fig. 1B).

Establishment of three new cryptic species based on molecular data and male genitalia
Eighteen of 22 morphological species were well supported by the ABGD result.M. angusta Wang, Li, Wang &Che, 2014 andM. mckittrickae Wang, Che &Wang, 2009 were considered as one MOTU.He et al. (2021) found some stable morphological differences between the two species, although the genetic distance between them was only ~ 5%.M. spinifera Bey-Bienko, 1958, M. bisignata Bey-Bienko, 1970, and M. paratransversa He & Wang, 2021 were all divided into two MO-TUs.These results suggest that it was insufficient and challenging to distinguish the specimens of Margattea only based on the external morphological characters.Therefore, we examined the male genitalia of M. spinifera, M. bisignata, and M. paratransversa carefully.For M. spinifera, the left end of the accessory sclerite of samples CY_7 and SP9 is trigonate (Fig. 2B), while that of samples CY_1, CY_2, CY_3, CY_4, CY_6, KY349628, MW970274, KY349636, KY349639 and MW970278 is expanded with fuzz (Fig. 2A).For M. bisignata, the left phallomere of samples SY_2 and SP4 has a short spiny process (Fig. 2D), while that of samples SY_1, SY_3, SY_4, SY_5, SY_6, SY_7, SY_8, SY_9, SY_10, MW970310, MW970317, MW970315, KY349596, KY349607, KY349603 and KY349604 had a long spine process (Fig. 2C).For M. paratransversa, the apex of median phallomere of samples HD_3 and SHD_1 is enlarged and forceps-shaped (Fig. 2F), while that of samples MW970262 and MW970263 has a slightly curved spine (Fig. 2E).Diagnosis.Body small, usually yellowish brown.Interocellar distance slightly wider than the distance between eyes, narrower than the distance between antennal sockets.The fifth maxillary palp expanded, the third and fourth palpi both longer than the fifth palp.Pronotum subelliptical, broader than long, the disc usually with symmetrical maculae and stripes.Tegmina and hind wings fully developed, mostly both extending beyond the end of abdomen.Tegmina with M and CuA radial, M straight with 4-7 branches.Hind wings with ScP and RA expanded at apex, CuA with 4-6 complete branches.Anteroventral margin of front femur Type B 2 or B 3 .Four proximal tarsomeres with pulvilli.The pretarsi with arolium, tarsal claws symmetrical and specialized, with minute denticles on ventral margins.Eighth abdominal tergum usually specialized, with a tuft of setae in the middle near posterior margin.Supra-anal plate usually short and transverse, paraprocts similar and flaky.Cerci long, with setae on the ventral surface.Male subgenital plate symmetrical or slightly asymmetrical.Styli symmetrical and cylindrical, rarely asymmetrical or non-cylindrical.Male genitalia.
Left phallomere small, irregularly bone-shaped, mostly with spine-like process.
Median phallomere slender, rod-shaped, the apex irregular and variable; accessory sclerite complicated, generally arched.Hook phallomere on right side, apex usually curved inwards.Differential diagnosis.The genus Margattea is supposedly closely related to Chorisoserrata (Wang et al. 2023) and morphologically similar to Balta, but Margattea could be distinguished from Chorisoserrata and Balta by the following characteristics.The genus Margattea can be distinguished from Balta (Asia, Africa, and parts of Oceania) by the following characteristics: 1) anteroventral margin of front femur Type B 2 or B 3 , in contrast to C 2 (but rarely B 3 ) in Balta; 2) the tarsal claws symmetrical and specialized, but in the latter, the tarsal claws asymmetrical and unspecialized.
The genus Margattea can be distinguished from Chorisoserrata (parts of Asia and Indonesia) by the following characteristics: 1) anteroventral margin
Description.Male.Coloration.Body pale yellow (Fig. 3A, B).Head yellowish brown.Face pale yellow.Interocular space with a wide brown transverse band.Ocellar spots white, interocellar space with a brown band.Antennal base pale yellowish brown, other segments dark brown (Fig. 3E).Maxillary palpi pale brown (Fig. 3J).Pronotal disc pale yellowish brown with reddish tan stripes, and two lateral borders pale linen-colored and transparent (Fig. 3F).Tegmina pale yellowish brown, hind wings brownish grey (Fig. 3G, H).Legs faint yellow.Abdomen yellowish brown, with black stripes along lateral margins of sterna and reaching the end of abdomen; both sides of each abdominal sternum with one small black spot on the inside of the longitudinal lines (Fig. 3B).Cerci yellowish brown to pale brown (Fig. 3L).Styli yellowish white (Fig. 3N).
Head.Vertex slightly exposed, interocellar distance slightly wider than the distance between eyes, narrower than the distance between antennal sockets (Fig. 3F).Fifth maxillary palpus expanded, third and fourth maxillary palpi nearly equal in length, both longer than fifth maxillary palpus (Fig. 3J).Pronotum subelliptical, broader than long, anterior and posterior margins nearly straight, and postero-lateral angle blunt and round; disc with symmetrical spots and stripes (Fig. 3E).Tegmina and hind wings.Tegmina and hind wings fully developed, both extending beyond the end of abdomen (Fig. 3A, B).Tegmina with M and CuA radial, M straight with three complete branches and one incomplete branch.Hind wings with ScP and RA expanded at apex, M simple, without branches; CuA with four complete branches (Fig. 3G, H).Legs.Anteroventral margin of front femur Type B 3 (Fig. 3K).Four proximal tarsomeres with pulvilli.The pretarsi with arolium, tarsal claws symmetrical and specialized, with minute denticles on ventral margins.
Diagnosis.This species is similar to M. speciosa Liu & Zhou, 2011 in general appearance, but can be differentiated from the latter by the following characters: 1) styli foot-shaped, while in the latter conical; 2) left phallomere with a long, curved spine, absent in the latter; and 3) accessory sclerite I without a brush-like structure at apex, while in the latter, accessory sclerite I with a brushlike structure at apex.
Etymology.The specific epithet is derived from the Latin word pedatus, referring to the foot-shaped styli.
Head.Vertex slightly exposed, interocellar distance slightly much wider than the distance between eyes, narrower than the distance between antennal sockets (Fig. 4E).Pronotum subelliptical, broader than long, anterior and posterior margins nearly straight, and postero-lateral angle blunt and round; disc with symmetrical but irregular spots and maculae (Fig. 4F).Fifth maxillary palpus expanded, third and fourth maxillary palpi both longer than fifth maxillary palpus (Fig. 4J).Tegmina and hind wings.Tegmina and hind wings fully developed, both extending beyond the end of abdomen (Fig. 4A, B).Tegmina M and CuA radial, M straight with six complete branches.Hind wings with ScP and RA expanded at apex, M simple, without branches; CuA with four complete branches (Fig. 4 G, H).Legs.Anteroventral margin of front femur Type B 2 (Fig. 4K).Four proximal tarsomeres with pulvilli.The pretarsi with arolium, tarsal claws symmetrical and specialized, with minute denticles on ventral margins.
Abdomen and genitalia.Eighth abdominal tergum specialized, with a tuft of bristles in the middle (Fig. 4I).Supra-anal plate symmetrical, anterior margin straight and truncate, the middle of posterior margin slightly concave.Paraprocts simple, similar, and flaky.Cerci long, with setae on the ventral surface (Fig. 4L).Subgenital plate asymmetrical.Styli dissimilar and spherical, the left stylus significantly smaller than the right stylus (Fig. 4N).Left phallomere small, irregular bone-shaped, with a slender curved spine (Fig. 4M).Median phallomere slender rod-shaped, with a curved spine at apex; accessory sclerite I arched, left end expanded, right end with a cluster of thorns; accessory sclerite II brush-shaped; accessory sclerite III slender rod-shaped (Fig. 4N).Hook phallomere on the right side, apex curved hook-shaped (Fig. 4O).
Female.Similar to the male.Subgenital plate symmetrical, middle posterior margin concave inward (Fig. 4C, D).Diagnosis.This species is similar to M. concava Wang, Che & Wang, 2009 in general appearance, but can be differentiated from the latter by the following characters: 1) styli dissimilar and spherical, the left stylus significantly smaller than the right stylus; while in the latter, styli similar and conical; 2) left phallomere with a slender curved spine, absent in the latter.
Etymology.The specific name is derived from the Latin words, bi and sphaericus, referring to the dissimilar and spherical styli.

Margattea undulata
Description.Male.Coloration.Body, head and face yellowish brown (Fig. 5A,  B).Interocular space with a brown transverse band.Ocellar spots small, yellowish white.Antennal base yellowish brown, other segments black-brown.The third and fourth maxillary palpi yellowish brown, the fifth palpus maxillary blackish brown (Fig. 5J).Pronotal disc yellowish brown with reddish tan spots and stripes, and two lateral borders pale linen-colored and transparent (Fig. 5E).Legs yellowish brown, with black spots at the base of the tibial spines.Tegmina yellowish brown, hind wings brownish grey (Fig. 5G, H).Abdomen yellowish brown, both sides of each abdominal sternum with one small round black spot on the inside of the longitudinal lines.Cerci yellowish brown (Fig. 5L).Styli pale yellow (Fig. 5N).
Head.Vertex slightly exposed, interocellar distance slightly wider than the distance between eyes, narrower than the distance between antennal sockets (Fig. 5F).Pronotum subelliptical, broader than long, anterior and posterior margins nearly straight, and postero-lateral margin blunt and round; disc with symmetrical spots and stripes (Fig. 5E).Fifth maxillary palpus expanded, third and fourth maxillary palpi nearly equal in length, both twice as long as fifth maxillary palpus (Fig. 5J).Tegmina and hind wings.Tegmina and hind wings fully developed, both extending beyond the end of abdomen (Fig. 5A, B).Tegmina with M and CuA radial, M straight with seven complete branches.Hind wings with ScP and RA expanded at apex, M simple, without branches; CuA with four complete branches (Fig. 5G, H).Legs.Anteroventral margin of front femur Type B 3 (Fig. 5K).Four proximal tarsomeres with pulvilli.The pretarsi with arolium, tarsal claws symmetrical and specialized, with minute denticles on ventral margins.Abdomen and genitalia.Eighth abdominal tergum specialized, with a tuft of setae near the dis- tinctly concave middle posterior margin (Fig. 5I).Supra-anal plate symmetrical, anterior margin straight and truncate, posterior margin obtusely round.Paraprocts simple, similar and flaky.Cerci long, with setae on the ventral surface (Fig. 5L) Subgenital plate nearly symmetrical, anterior margin distinctly concave in the middle.Styli similar, slender; interstylar margin sinuate, left side with five or six small spines, right side with 5-7 small spines (Fig. 5N).Left phallomere small, irregular bone-shaped, with a small spine (Fig. 5M).Median phallomere slender rod-shaped, with a bristle brush at apex; accessory sclerite I arched, two ends enlarged, right end with a row of spines; accessory sclerite II with three lamellar structures with small spines; accessory sclerite III sickle-shaped (Fig. 5N).Hook phallomere on the right side, apex curved inwards with a short spine (Fig. 5O).
Female.Similar to the male but body and wings somewhat shorter (Fig. 5C, D).
Diagnosis.This species is similar to M. flexa Wang et al., 2014 in general appearance, but can be differentiated from the latter by the following characters: 1) interstylar margin sinuate, left side with 4-6 small spines, right side with 4-7 small spines; while in the latter, interstylar margin strongly produced, whose lateral sides upturned and scattered with spines; 2) left phallomere irregular bone-shaped, without a small spine; while in the latter, left phallomere irregular bone-shaped, with two spines; 3) accessory sclerite II with three lamellar structures with small spines; while in the latter, accessory sclerite II with lamellar structure without small spines.
Etymology.The specific name is derived from the Latin word undulatus, which refers to the sinuate interstylar margin.

Margattea semicircularis
Description.Male.Coloration.Body, head and face yellowish brown (Fig. 6A,  B).Interocular space with a wide brown transverse band.Ocellar spots small, white, with brown spots beside them.Antennal base pale yellowish brown, other segments brown (Fig. 6D).Maxillary palpi dark brown (Fig. 6H).Pronotal disc yellowish brown with reddish tan spots and stripes, and two lateral borders pale linen-colored and transparent (Fig. 6C).Legs yellowish brown, with black spots at the base of the tibial spines.Tegmina pale yellowish brown, hind wings brownish grey (Fig. 6F, G).Abdomen yellowish brown, with black stripes along lateral margins of sterna and reaching the end of abdomen; both sides of each abdominal sternum with one small black spot on the inside of the longitudinal lines (Fig. 6B).Cerci yellowish brown (Fig. 6K).Styli pale yellow (Fig. 6M).
Head.Vertex slightly exposed, interocellar distance wider than the distance between eyes, narrower than the distance between antennal sockets (Fig. 6C, D).Pronotum subelliptical, broader than long, anterior and posterior margins nearly straight, and postero-lateral angle blunt and round; disc with symmetrical spots and stripes (Fig. 6C).Maxillary palpi slender, fifth maxillary palpus expanded, third and fourth maxillary palpi nearly equal in length, both longer than fifth maxillary palpus (Fig. 6H).Tegmina and hind wings.Tegmina and hind wings fully developed, both extending beyond the end of abdomen (Fig. 6A, B).Tegmina with M and CuA radial, M straight with seven complete branches.Hind wings with ScP and RA expanded at apex; M simple, without branches; CuA with five complete branches (Fig. 6F, G).Legs.Anteroventral margin of front femur Type B 3 (Fig. 6I).Four proximal tarsomeres with pulvilli.The pretarsi with arolium, trsal claws symmetrical and specialized, with minute denticles on ventral margins.
Abdomen and genitalia.Eighth abdominal tergum specialized, with a heartshaped transparent area and a tuft of bristles in the middle (Fig. 6E).Supra-anal plate symmetrical, anterior margin straight and truncate, the middle of posterior margin slightly produced.Paraprocts simple, similar, and flaky.Cerci long, setae on the ventral surface (Fig. 6K).Subgenital plate nearly symmetrical, anterior margin slightly concave in the middle.Styli similar, conical; interstylar margin strongly semicircular produced, both sides with spines (Fig. 6K).Left phallomere small, irregular, bone-shaped, with two small spines (Fig. 6J).Median phallomere curved hook rod-shaped, with a row of spines at apex.Accessory sclerite I arched, two ends enlarged, left end trigonate, right end with a row of spines; accessory sclerite II long transverse (Fig. 6L, M).Hook phallomere on the right side, apex curved inwards with a short spine (Fig. 6N).
Diagnosis.This species is similar to M. spinifera Bey-Bienko, 1958a in general appearance, but can be differentiated from the latter by the following characters: 1) left phallomere with two small spines; while the latter, left phallomere with three spine-like processes; 2) accessory sclerite I arched, left end trigonate; while in the latter, accessory sclerite I arched, left end expanded with fuzz; 3) accessory sclerite II long transverse, and with two lamellar structures with a row of spines, while in the latter, without other accessory sclerites.
Etymology.The scientific name is derived from the Latin word semicircularis, which indicates the interstylar margin has a semicircular protrusion.

Margattea parabisignata
Head.Vertex slightly exposed, interocellar distance wider than the distance between eyes, narrower than the distance between antennal sockets (Fig. 7F).Pronotum subelliptical, broader than long, anterior and posterior margins nearly straight, and postero-lateral angle blunt and round; disc with symmetrical spots but without stripes (Fig. 7E).Fifth maxillary palpus expanded, third and fourth maxillary palpi nearly equal in length, both longer than fifth maxillary palpus (Fig. 7J).Tegmina and hind wings.Tegmina and hind wings fully developed, both extending beyond the end of abdomen (Fig. 7A, B).Tegmina with M and CuA radial, M straight with seven complete branches.Hind wings with ScP and RA expanded at apex, M simple, without branches; CuA with four complete branches (Fig. 7G, H).Legs.Anteroventral margin of front femur Type B 2 (Fig. 7K).Four proximal tarsomeres with pulvilli.The pretarsi with arolium, tarsal claws symmetrical and slightly specialized, with minute denticles on ventral margins.
Abdomen and genitalia.Eighth abdominal tergum specialized, with a sparse tuft of bristles in the middle (Fig. 7I).Supra-anal plate symmetrical, the middle of anterior margin slightly concave, posterior margin arcuate produced with setae.Paraprocts simple, similar and flaky, obtuse at apex and each with a spiniform process at the base (Fig. 7L).Subgenital plate nearly symmetrical, anterior margin distinctly concave in the middle, left and right margins both produced in the middle, posterior margin truncate.Styli similar, conical.Cerci long, with setae on the ventral surface (Fig. 7N).Left phallomere large, irregular, bone-shaped, and with spines processes, apex curved upwards with rod-like structure (Fig. 7M).Median phallomere slender rod-shaped, apex with sparse brush-like structure composed of spines of various sizes; accessory sclerite I arched; accessory sclerite II with a lamellar structure with small spines (Fig. 7N).Hook phallomere on the right side, apex curved inwards with a short spine (Fig. 7O).
Diagnosis.This species is similar to M. bisignata Bey-Bienko, 1970 in general appearance, but can be differentiated from the latter by the following characters: 1) left phallomere with a short spiny process; the latter with a long spine process; 2) median phallomere apex with sparse brush-like structure composed of spines of varying sizes; while in the latter, median phallomere curved at apex, sheet-like, and with brush-shaped structure.
Description.Male.Coloration.Body, head and face yellowish brown (Fig. 8A,  B).Interocular space with a wider brown transverse band.Ocellar spots big and white (Fig. 8F).Antennal base pale yellow, other segments yellowish brown to brown.Maxillary palpi yellowish brown (Fig. 8J).Pronotal disc yellowish brown with dark brown spots and maculae, and two lateral borders pale linen-colored and transparent (Fig. 8E).Legs yellowish brown, with black spots at the base of the tibial spines.Tegmina pale yellowish brown, hind wings transparent, brownish grey (Fig. 8G, H).Abdomen yellowish brown, with black stripes along lateral margins of sterna and reaching the end of abdomen; both sides of each abdominal sternum with one small round black spot on the inside of the longitudinal lines.Cerci yellowish brown (Fig. 8L).Styli yellowish white (Fig. 8N).
Head.Vertex slightly exposed, interocellar distance wider than the distance between eyes, narrower than the distance between antennal sockets (Fig. 8F).Pronotum subelliptical, broader than long, anterior and posterior margins nearly straight, and postero-lateral angle blunt and round; disc with symmetrical irregular spots and maculae (Fig. 8E).Fifth maxillary palpus expanded, third and fourth maxillary palpi both longer than fifth maxillary palpus (Fig. 8J).Tegmina and hind wings.Tegmina and hind wings fully developed, both extending beyond the end of abdomen (Fig. 8A, B).Tegmina with M and CuA radial, M straight with six complete branches.Hind wings with ScP and RA expanded at apex, M simple, without branches; CuA with four complete branches (Fig. 8G,  H).Legs.Anteroventral margin of front femur Type B 2 (Fig. 8K).Four proximal tarsomeres with pulvilli.The pretarsi with arolium, tarsal claws symmetrical and specialized, with minute denticles on ventral margins.
Abdomen and genitalia.Eighth abdominal tergum specialized, with a tuft of bristles in the middle (Fig. 8I).Supra-anal plate symmetrical, anterior margin straight and truncate, posterior margin produced, slightly concave in the middle.Paraprocts simple, similar, and flaky.Cerci long, setae on the ventral surface (Fig. 8L).Subgenital plate nearly symmetrical, anterior margin slightly concave in the middle.Styli similar, slender, conical; interstylar margin irregular produced (Fig. 8N).Left phallomere complex, irregular bone-shaped, with a long spine and three small spines (Fig. 8M).Median phallomere slender rodshaped, with a forceps-shaped apex.Accessory sclerite I arched; accessory sclerite II with a transverse rod with denticulate (Fig. 8N).Hook phallomere on the right side, apex slightly curved inwards with a short spine (Fig. 8O).Female.Similar to the male.Diagnosis.This species is similar to M. transversa He & Wang, 2021 in general appearance, but can be differentiated from the latter by the following characters: 1) left phallomere with a long spine; the latter with three long spine-like processes; 2) median phallomere with a forceps-shaped apex; while in the latter, median phallomere apex with a curved spine.
Etymology.The specific name forcipatus, derived from Latin, refers to the median phallomere with a forceps-shaped apex.
In this study, we initially determined three morphospecies, namely "M.spinifera", "M.bisignata", and "M.paratransversa", whose individuals are almost indistinguishable.In contrast, these morphospecies are each divided into two MO-TUs in molecular species delimitation.We hence examined the male genitalia of different samples from each of these morphospecies and found differences in the accessory sclerite I of "M.spinifera" (Fig. 2A, B), the left phallomere of "M.bisignata" (Fig. 2C, D), and the median phallomere of "M.paratransversa" (Fig. 2E, F).With the assistance of male genitalia examination, these MOTUs were determined as different species.This also occurs in other genera in Blattodea, where large genetic distances among closely related species might occur despite small differences in external morphology (Bai et al. 2018;Han et al. 2022;Zhu et al. 2022).Nine Cryptocercus species were extremely similar in external morphology, five of which could be distinguished according to chromosome number and female genital characteristics.The other four species could not be distinguished solely based on chromosome number and female genital characteristics, but they could be distinguished by combining these with molecular species definition (Bai et al. 2018).Anaplecta omei Bey-Bienko, 1958b could be distinguished from the other three species with very similar external morphology by molecular species definition and female genitalia characteristics (Zhu et al. 2022).Pseudoeupolyphaga simila (Qui, 2022) was extremely similar to Pseudoeupolyphaga yunnanensis (Chopard, 1922) in external morphology, but they could be distinguished by combining female genitalia characteristics, oothecae characteristics, and molecular species definition (Han et al. 2022).In Blattodea and even the insect community, there is an increasing occurrence of closely resembling morphologies that do not necessarily belong to the same species.It is no longer possible to determine species only by morphological characteristics; it is also necessary to recognize species from various aspects, e.g., endosymbionts, cytological characteristics, and ecological characteristics.

Figure 1 .
Figure 1.Maximum likelihood (ML) tree and species delimitation of Margattea based on COI sequence.Branches labels are bootstrap support percentage.Colored bars indicate different species delimitation by different methods A morphology (pink) B ABGD results (blue).
In conclusion, three cryptic new species, M. parabisignata Li & Che, sp.nov., M. semicircularis Li & Che, sp.nov., and M. forcipata Li & Che, sp.nov., are discovered mainly based on the male genitalia with the help of the molecular data.

Figure 3 .
Figure 3. Margattea pedata Li & Che, sp.nov.A, B, E-O male C, D female A holotype, dorsal view B holotype, ventral view C paratype, dorsal view D paratype, ventral view E pronotum, dorsal view F head, ventral view G tegmen, ventral view H hind wing, ventral view I eighth abdominal tergum, ventral view J maxillary palpi segments 3-5 K front femur, ventral view L supra-anal plate, ventral view M left phallomere, ventral view N subgenital plate and median phallomere, ventral view O hook phallomere, ventral view.Scale bars: 5 mm (A-D, G, H); 1 mm (E, F, I-L, N); 0.5 mm (M, O).

Figure 4 .
Figure 4. Margattea bisphaerica Li & Che, sp.nov.A, B, E-O male C, D female A holotype, dorsal view B holotype, ventral view C paratype, dorsal view D paratype, ventral view E pronotum, dorsal view F head, ventral view G tegmen, ventral view H hind wing, ventral view I eighth abdominal tergum, ventral view J maxillary palpi segments 3-5 K front femur, ventral view L supra-anal plate, ventral view M left phallomere, ventral view N subgenital plate and median phallomere, ventral view O hook phallomere, ventral view.Scale bars: 5 mm (A-D, G, H); 1 mm (E, F, I-K, N); 0.5 mm (L, M, O).

Figure 5 .
Figure 5. Margattea undulata Li & Che, sp.nov.A, B, E-O male C, D female A holotype, dorsal view B holotype, ventral view C paratype, dorsal view D paratype, ventral view E pronotum, dorsal view F head, ventral view G tegmen H hind wing I eighth abdominal tergum, ventral view J maxillary palpi segments 3-5 K front femur, ventral view L supra-anal plate, ventral view M left phallomere, ventral view N subgenital plate and median phallomere, ventral view O hook phallomere, ventral view.Scale bars: 5 mm (A-D, G, H); 1 mm (E, F, I-L, N); 0.5 mm (M, O).

Figure 7 .
Figure 7. Margattea parabisignata Li & Che, sp.nov.A, B, E-O male C, D female A holotype, dorsal view B holotype, ventral view C paratype, dorsal view D paratype, ventral view E pronotum, dorsal view F head, ventral view G tegmen, ventral view H hind wing, ventral view I eighth abdominal tergum, ventral view J maxillary palpi segments 3-5 K front femur, ventral view L supra-anal plate, ventral view M left phallomere, ventral view N subgenital plate and median phallomere, ventral view O hook phallomere, ventral view.Scale bars: 5 mm (A-D, G, H); 1 mm (E, F, I-L, N); 0.5 mm (M, O).

Figure 8 .
Figure 8. Margattea forcipata Li & Che, sp.nov.A, B, E-O male C, D female A holotype, dorsal view B holotype, ventral view C paratype, dorsal view D paratype, ventral view E pronotum, dorsal view F head, ventral view G tegmen, ventral view H hind wing, ventral view I eighth abdominal tergum, ventral view J maxillary palpi segments 3-5 K front femur, ventral view L supra-anal plate, ventral view M left phallomere, ventral view N subgenital plate and median phallomere, ventral view O hook phallomere, ventral view.Scale bars: 5 mm (A-D, G, H); 1 mm (E, F, I-L, N); 0.5 mm (M, O).

Table 1 .
Primers used in the molecular study.

Table 2 .
Samples used in the molecular study.