Phylogeny and Integrative Taxonomy of the Genera Gymnaetoides and Pseudotachycines (Orthoptera: Rhaphidophoridae)

Simple Summary The genera Gymnaetoides and Pseudotachycines are morphologically homogeneous and species could not be classified according to the given generic diagnosis. Here, we reconstruct the phylogeny, and the results show that both Gymnaetoides and Pseudotachycines are paraphyletic. Therefore, we revise their taxonomy based on morphological characters and molecular data. A new genus is erected, and six new combinations and 15 new species are proposed. Moreover, we find that the shape of the male genitalia is important character for identifying these genera. Abstract The genera Gymnaetoides and Pseudotachycines are endemic to China and are morphologically homogeneous. The few available diagnostic characters make species identification particularly challenging. Species cannot be classified according to the given generic diagnosis, and phylogenetic analyses have not been reported. Here, we reconstruct the phylogeny using Bayesian inference and maximum likelihood and employ four approaches to delimit species. The results suggest that both Gymnaetoides and Pseudotachycines are paraphyletic. Therefore, we revise their taxonomy based on the combination of morphological characters and molecular data. A new genus Homotachycines Zhu & Shi gen. nov. is erected, and six new combinations are proposed. Species delimitation identifies 15 new species and one new subspecies: Gymnaetoides huangshanensis, G. petalus, G. yangmingensis, G. lushanensis, Pseudotachycines procerus, P. procerus guizhouensis, P. zhengi, P. nephrus, P. sagittus, P. fengyangshanensis, Homotachycines triangulus, H. quadratus, H. baokangensis, H. fusus, H. concavus, and H. qinlingensis sp. nov. Moreover, we find that the shapes of the dorsal lateral lobes and the dorsal median lobe of the male genitalia are also important characters for identifying these genera and that the shapes of the dorsal and lateral sclerites of the male genitalia are suitable for the classifications of species.


Introduction
There are nine extant subfamilies and one extinct subfamily of Rhaphidophoridae, of which two subfamilies (Aemodogryllinae and Rhaphidophorinae) are recorded from China [1]. The subfamily Aemodogryllinae includes two tribes Aemodogryllini and Diestramimini, while Rhaphidophorinae includes one tribe Rhaphidophorini. Cave crickets prefer dark and moist habitats, most of them usually hide under litter or loose bark during the day and come out to hunt or mate at night, while others adapt to cave life [2][3][4]. They are wingless and do not have tibial tympana or a stridulatory organ. Moreover, their PP > 0.95 or BS > 70 are given at the nodes. In ML tree, PT868HB is recovered as sister to PT746HB (BS = 92) and together they are sister to PT819HB, PT820HB, and PT867HB (BS = 100). Results of molecular species delimitation based on ABGD, jMOTU, bPTP, GMYC, and morphological species are annotated on the right side of the tree.

Species Delimitation
The results of the four species delimitation methods were inconsistent (Figure 1). Although the ABGD and jMOTU analyses were consistent with the morphological species in clustering 22 MOTUs, the bPTP analysis generated 29 MOTUs and the GMYC analysis resulted in 25 MOTUs. Moreover, there was also inconsistency between molecular data  According to the given generic diagnosis, Gymnaetoides is characterized by the distinctly sclerotized apex of the dorsolateral lobes of the male genitalia and the normally developed male epiproct and paraproct, while Pseudotachycines is characterized by the sclerotized apex of the dorsolateral lobes of the male genitalia and the distinctly prolonged male paraproct. Therefore, the clade Gymnaetoides contains seven species and P. deformis and P. yueyangensis should be transferred to it. In the clade Pseudotachycines, only the known species P. volutus and the five new species are included. The third clade represents a new taxon and thus a new genus Homotachycines Zhu & Shi gen. nov. is proposed.
Homotachycines is sister to Pseudotachycines, and they are together sisters to Gymnaetoides. Moreover, P. ovalilobatus and G. acutilobatus should be transferred to Homotachycines.

Species Delimitation
The results of the four species delimitation methods were inconsistent ( Figure 1). Although the ABGD and jMOTU analyses were consistent with the morphological species in clustering 22 MOTUs, the bPTP analysis generated 29 MOTUs and the GMYC analysis resulted in 25 MOTUs. Moreover, there was also inconsistency between molecular data and morphological species. In Pseudotachycines procerus, specimens from Guizhou and Anhui formed two MOTUs in all of the four species delimitation methods. Despite the inconsistency of different species delimitation methods or the inconsistency of morphological and genetic data, most species (17 of 22) can successfully be identified.

1.
The apical area of the dorsal median lobe of the male genitalia divides into two lobes; the dorsal lateral lobe of the male genitalia is of nearly equal length as the dorsal median lobe, in which the apical area is distinctly sclerotized . . . . . . . . . Gymnaetoides

Distribution. China (Zhejiang).
Remarks. The species was recorded in Zhejiang (Gutian Mountain, Tianmu Mountain) and Anhui (Huang Mountain). In the original description, 65-68 inner and 64-69 outer spines on dorsal surface of the hind tibia were reported. However, after examining the type and topotype specimens, 30-39 inner and 37-40 outer spines on dorsal surface of the hind tibia in specimens from Zhejiang were observed, while 60-69 inner spines and 64-69 outer spines were observed in specimens collected from Anhui. This was also indicated by Qin [16]. Moreover, all of the four species delimitation approaches suggest that the specimens from Zhejiang and Anhui are two separate species (Figure 1). Therefore, we consider the population from Anhui as a new species, Gymnaetoides huangshanensis Zhu & Shi sp. nov. (described below). G. testaceus is currently found only in Zhejiang and the description of spines on hind tibia should be changed to "hind tibia with 30-39 inner spines and 37-40 outer spines on dorsal surface". In addition, the line drawings of the male genitalia did not adequately represent the characters (Figures 3 and 4 in [13]). The photograph of type specimen is shown in Figure 2A.
Gymnaetoides surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and one outer spine on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur unarmed on ventral surface, internal and external genicular lobes unarmed; tibia with 60-69 inner spines and 64-68 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine shorter than hind basitarsus. Hind basitarsus with one dorsal apical spine.   Posterior margins of all abdominal tergites straight. Epiproct ligulate, paraproct triangular in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia quadrangular, basal area broad, with concavity, apical area slightly concave; basal area of dorsal median lobe with paired lateral sclerites, nearly semicircular, apical area divided into two lobes; dorsal lateral lobes broad, nearly equal in length to dorsal median lobe, apical area sclerotized, spinous; ventral lateral lobes tiny, much shorter than dorsal lateral lobes; ventral median lobe short. Subgenital plate quadrangular, transverse and wide, posterior margin slightly concave. Remarks. This species was designated as the type species of Pseudotachycines, which was characterized by the prolonged male paraproct [14]. However, after examining the type specimen, we did not find the specialized paraproct. Moreover, the line drawings of the male genitalia did not adequately represent the characters (Figures 4 and 5 in [14]). The photograph of the type specimen is shown in Figure 2B. The result of phylogenetic analyses also clusters this species in genus Gymnaetoides ( Figure 1). It should be transferred into the genus Gymnaetoides according to the characters of the male genitalia: the apical area of the dorsal median lobe divided into two lobes, the dorsal lateral lobes broad, nearly equal in length to dorsal median lobe, the apical area sclerotized. Distribution. China (Hubei). Remarks. The species was classified into Pseudotachycines based on the prolonged male paraproct [14]. However, after examining the type specimen, this feature was not distinct. The line drawings of the male genitalia did not adequately represent the characters (Figures 8 and 9 in [14]). The photograph of the type specimen is shown in Figure 2C. The dorsal sclerite of the male genitalia is rectangular, much longer than wide; the lateral sclerite of the dorsal median lobe is nearly rectangular, with the apical area divided into two lobes; the dorsal lateral lobe is broad, nearly equal in length to the dorsal median lobe, with the apical area sclerotized; the ventral lateral lobe and the ventral median lobe are short. According to the characters of the male genitalia, it should be transferred into the genus Gymnaetoides.
Gymnaetoides Remarks. After examining the type specimen, we found that the line drawings of the male genitalia did not adequately represent the characters (Figures 10 and 11 in [14]). The photograph of the type specimen is shown in Figure 2D. The apical area of the dorsal median lobe of the male genitalia divides into two lobes; the dorsal lateral lobe is broad, nearly equal in length to the dorsal median lobe, with the apical area sclerotized. According to the characters of the male genitalia and the phylogenetic analyses, we transfer it into genus Gymnaetoides.
Gymnaetoides    , but it differs from the latter one by the shape of the male genitalia. The dorsal sclerite of the male genitalia is quadrangular, with a concavity on the basal area; the lateral sclerite of the dorsal median lobe is semicircular; the apical area of the dorsal lateral lobe is spinous.
Pronotum long, anterior margin of disc straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe with one small spine, external genicular lobe with one long spine; tibia with two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and one outer spine on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur unarmed or with one spine on ventral surface, internal genicular lobe with one small spine; tibia with 50-56 inner spines and 51-53 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two Etymology. The name of the new species derives from the Greek word 'petal' (petaloid), referring to the petaloid dorsal sclerite of the male genitalia.
Diagnosis. The new species can easily be distinguished from known congeneric species by the shape of the male genitalia: the dorsal sclerite of the male genitalia is petaloid and the lateral sclerite of the dorsal median lobe is slender.
Anterior margin of pronotum straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe with one small spine, external genicular lobe with one long spine; tibia with two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and one outer spine on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur unarmed on ventral surface; tibia with 87-92 inner spines and 84-86 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine shorter than hind basitarsus. Hind basitarsus with one dorsal apical spine.
All abdominal tergites without process. Epiproct ligulate, paraproct slightly longer than epiproct. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia petaloid; basal area of dorsal median lobe with paired lateral sclerites, slender, apical area divided into two lobes; dorsal lateral lobes broad, nearly equal in length to dorsal median lobe, apical area sclerotized, spinous; ventral lateral lobes cylindrical, much shorter than dorsal lateral lobes; ventral median lobe short. Subgenital plate transverse and wide, posterior margin widely concave.
Female  All abdominal tergites without process. Epiproct ligulate, paraproct triangular in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia quadrangular, basal area wide, with concavity; basal area of dorsal median lobe with paired lateral sclerites, semicircular, apical area divided into two lobes; dorsal lateral lobes broad, nearly equal in length to dorsal median lobe, apical area spinous; ventral lateral lobes much shorter than dorsal lateral lobes; ventral median lobe short. Subgenital plate transverse and wide, posterior margin straight.
Female: Appearance is similar to male. Ovipositor longer than half the length of hind femur, slightly curved upward, dorsal margin smooth, apical area of ventral margin denticulate. Subgenital plate trapezoidal, posterior margin slightly protruding.
Coloration  Etymology. The name of the new species refers to the type locality. Diagnosis. The new species is similar to Gymnaetoides yueyangensis (Qin, Liu & Li, 2017), but it differs from the latter one by the shape of the male genitalia. The dorsal sclerite of the male genitalia is quadrangular, with a concavity on the basal area; the lateral sclerite of the dorsal median lobe is semicircular; the apical area of the dorsal lateral lobe is spinous.
Pronotum long, anterior margin of disc straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe with one small spine, external genicular lobe with one long spine; tibia with two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and one outer spine on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur unarmed or with one spine on ventral surface, internal genicular lobe with one small spine; tibia with 50-56 inner spines and 51-53 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine nearly equal in length to hind basitarsus. Hind basitarsus with one dorsal apical spine.
All abdominal tergites without process. Epiproct ligulate, paraproct triangular in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia quadrangular, basal area wide, with concavity; basal area of dorsal median lobe with paired lateral sclerites, semicircular, apical area divided into two lobes; dorsal lateral lobes broad, nearly equal in length to dorsal median lobe, apical area spinous; ventral lateral lobes much shorter than dorsal lateral lobes; ventral median lobe short. Subgenital plate transverse and wide, posterior margin straight.
Female: Appearance is similar to male. Ovipositor longer than half the length of hind femur, slightly curved upward, dorsal margin smooth, apical area of ventral margin denticulate. Subgenital plate trapezoidal, posterior margin slightly protruding.
Coloration Etymology. The name of the new species derives from the type locality. Diagnosis. The new species is similar to Gymnaetoides huangshanensis Zhu & Shi sp. nov., but it can be distinguished by the shapes of lateral sclerite of the dorsal median lobe and the dorsal lateral lobe of the male genitalia. The lateral sclerite of the dorsal median lobe is spoon shaped and the apical area of the dorsal lateral lobe is slightly sclerotized.
Description. Male: Body medium sized in Gymnaetoides. Fastigium verticis with two conical tubercles, apices divided, directing forward. Eyes ovoid, protruding forward; median ocellus oval, located between antennal sockets; lateral ocelli circular, situated on lateral margins of basal fastigium verticis. Apical segment of maxillary palpus much longer than subapical segment, apex inflated, globular.  Diagnosis. Body is medium sized in Aemodogryllini. The male paraproct are specialized, slightly or much longer than epiproct. The male genitalia have unpaired dorsal sclerite, smaller than that of the other genera in Aemodogryllini; the dorsal median lobe of the male genitalia has paired lateral sclerites at the basal area, the apical area does not divide; the dorsal lateral lobes of the male genitalia are broad and folded, much longer than dorsal median lobe, with the apical area slightly sclerotized, almost as wide as the basal area.  Pronotum long, anterior margin of disc straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe with one small spine, external genicular lobe with one long spine; tibia with two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and two outer spines on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur unarmed on ventral surface; tibia with 65-75 inner spines and 63-71 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine slightly shorter than hind basitarsus. Hind basitarsus with one dorsal apical spine.
Posterior margins of all abdominal tergites straight. Epiproct ligulate, paraproct simple, triangular in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia quadrangular, basal area wide, with a concavity, apex slightly concave; basal area of dorsal median lobe with paired lateral sclerites, spoon-shaped, apical area divided into two lobes; dorsal lateral lobes broad, nearly equal in length to dorsal median lobe, apical area slightly sclerotized; ventral lateral lobes cylindrical, nearly equal in length to dorsal lateral lobes; ventral median lobe short. Subgenital plate transverse and wide, posterior margin straight.
Female: Appearance is similar to male. Ovipositor longer than half the length of hind femur, dorsal margin smooth, apical area of ventral margin denticulate. Subgenital plate triangular, apex acute.
Coloration. Body yellow. Pronotum and apex of abdomen yellowish brown. Diagnosis. Body is medium sized in Aemodogryllini. The male paraproct are specialized, slightly or much longer than epiproct. The male genitalia have unpaired dorsal sclerite, smaller than that of the other genera in Aemodogryllini; the dorsal median lobe of the male genitalia has paired lateral sclerites at the basal area, the apical area does not divide; the dorsal lateral lobes of the male genitalia are broad and folded, much longer than dorsal median lobe, with the apical area slightly sclerotized, almost as wide as the basal area. Remarks. Because the type species of genus Pseudotachycines is moved to genus Gymnaetoides, Pseudotachycines volutus is redesignated as the type species of the genus Pseudotachycines. After examining the type specimen, we found that the line drawings of the male genitalia did not adequately represent the characters (Figures 14-16 in [14]). The photographs of the type specimens are shown in Figure 8.  Remarks. Because the type species of genus Pseudotachycines is moved to genus Gymnaetoides, Pseudotachycines volutus is redesignated as the type species of the genus Pseudotachycines. After examining the type specimen, we found that the line drawings of the male genitalia did not adequately represent the characters (Figures 14-16 in [14]). The photographs of the type specimens are shown in Figure 8. Description. Male: Body medium sized in Pseudotachycines. Fastigium verticis with two conical tubercles, apices rounded, directing forward. Eyes ovoid, protruding forward; median ocellus oval, located between antennal sockets; lateral ocelli circular, situated on lateral margins of basal fastigium verticis. Apical segment of maxillary palpus much longer than subapical segment, apex inflated, globular.
Anterior margin of pronotum straight, posterior margin arcuate; lateral lobes longer 60 inner spines and 55-56 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine much longer than hind basitarsus. Hind basitarsus with one dorsal apical spine. All abdominal tergites without process. Epiproct ligulate, paraproct extended backward, slightly longer than epiproct, oblong in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia small and lunular; basal area of dorsal median lobe with paired lateral sclerites, oblong, apical area not divided; dorsal lateral lobes broad and folded, much longer than dorsal median lobe, subapical area with a process on dorsal surface, apical area slightly sclerotized, almost as wide as basal area; ventral lateral lobes and ventral median lobe short. Subgenital plate quadrangular, basal half broad, apical half narrow, posterior margin slightly concave.
Female: Appearance is similar to male. Ovipositor longer than half the length of hind femur, curved upward, dorsal margin smooth, apical area of ventral margin denticulate. Subgenital plate quadrangular, basal area broad, gradually narrowing to apex, posterior margin blunt.
Coloration. Dorsal surface of body light brown, ventral surface yellowish brown. Face with four longitudinal black stripes. Etymology. The name of the new species derives from the Latin word 'procer' ('process'), referring to the subapical area of the dorsal lateral lobe of the male genitalia with a process on the dorsal surface.
Diagnosis. The new species can easily be distinguished from known congeneric species by the shape of the male genitalia and paraproct. The dorsal sclerite of the male genitalia is lunular and the lateral sclerite of the dorsal median lobe is oblong. The subapical area of the dorsal lateral lobe of the male genitalia has a process on the dorsal surface. The male paraproct is oblong in the lateral view.
Anterior margin of pronotum straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe with one small spine, external genicular lobe with one long spine; tibia with two to three inner spines and two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and two outer spines on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur with three to five inner spines on ventral surface; tibia with 57-60 inner spines and 55-56 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine much longer than hind basitarsus. Hind basitarsus with one dorsal apical spine.
All abdominal tergites without process. Epiproct ligulate, paraproct extended backward, slightly longer than epiproct, oblong in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia small and lunular; basal area of dorsal median lobe with paired lateral sclerites, oblong, apical area not divided; dorsal lateral lobes broad and folded, much longer than dorsal median lobe, subapical area with a process on dorsal surface, apical area slightly sclerotized, almost as wide as basal area; ventral lateral lobes and ventral median lobe short. Subgenital plate quadrangular, basal half broad, apical half narrow, posterior margin slightly concave.
Female: Appearance is similar to male. Ovipositor longer than half the length of hind femur, curved upward, dorsal margin smooth, apical area of ventral margin denticulate. Subgenital plate quadrangular, basal area broad, gradually narrowing to apex, posterior margin blunt.
Coloration   Etymology. The name of the new species derives from the type locality. Diagnosis. The species is very similar to Pseudotachycines procerus Zhu & Shi sp. nov., the slight differences are as follows: the lateral sclerite of the dorsal median lobe of the male genitalia is semicircular, and the hind femur has one to three inner spines on the ventral surface, which is less than three to five inner spines of the latter one.
Pronotum long, anterior margin of disc straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe with one small spine, external genicular lobe with one long spine; tibia with two inner spines and two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and two outer spines on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur with one to three inner spines on ventral surface; tibia with 53-56 inner spines and 50-54 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine much longer than hind basitarsus. Hind basitarsus with one dorsal apical spine.
All abdominal tergites without process. Epiproct ligulate, paraproct extended backward, slightly longer than epiproct, oblong in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia small and lunular, basal area slightly concave; basal area of dorsal median lobe with paired lateral sclerites, semicircular, apical area not divided; dorsal lateral lobes broad and folded, much longer than dorsal median lobe, subapical area with a process on dorsal surface, apical area sclerotized, almost as wide as basal area; ventral lateral lobes and ventral median lobe short. Subgenital plate quadrangular, basal half broad, apical half narrow, posterior margin slightly concave.
Female: Unknown. Diagnosis. The new species can easily be distinguished from known congeneric species by the shape of the male genitalia and paraproct. The dorsal sclerite of the male genitalia is nearly semicircular, with a concavity at the basal area and the apical area broadly concave. The lateral sclerite of the dorsal median lobe of the male genitalia is oblong. The male paraproct is semicircular in the lateral view.
59 inner spines and 59-62 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine much longer than hind basitarsus. Hind basitarsus with one dorsal apical spine.
All abdominal tergites without process. Epiproct lingulate; paraproct slightly longer than epiproct, semicircular in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia nearly semicircular, basal area with a concavity, apical area broadly concave; basal area of dorsal median lobe with paired lateral sclerites, oblong, apical area not divided; dorsal lateral lobes broad and folded, much longer than dorsal median lobe, apical area weakly sclerotized, almost as wide as basal area; ventral median lobe and ventral lateral lobes short. Subgenital plate quadrangular, wide, posterior margin straight.  Description. Male: Body medium sized in Pseudotachycines. Fastigium verticis with two conical tubercles, apices rounded, directing forward. Eyes ovoid, protruding forward; median ocellus oval, located between antennal sockets; lateral ocelli circular, situated on lateral margins of basal fastigium verticis. Apical segment of maxillary palpus much longer than subapical segment, apex inflated, globular.
Pronotum long, anterior margin straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe with one small spine, external genicular lobe with one long spine; tibia with two inner spines and two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with 1 long spine; tibia with one inner spine and two to three outer spines on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur with two inner spines on ventral surface; tibia with 56-59 inner spines and 59-62 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine much longer than hind basitarsus. Hind basitarsus with one dorsal apical spine.
All abdominal tergites without process. Epiproct lingulate; paraproct slightly longer than epiproct, semicircular in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia nearly semicircular, basal area with a concavity, apical area broadly concave; basal area of dorsal median lobe with paired lateral sclerites, oblong, apical area not divided; dorsal lateral lobes broad and folded, much longer than dorsal median lobe, apical area weakly sclerotized, almost as wide as basal area; ventral median lobe and ventral lateral lobes short. Subgenital plate quadrangular, wide, posterior margin straight.
Female: Appearance is similar to male. Ovipositor longer than half the length of hind femur, dorsal margin smooth, apical area of ventral margin denticulate. Subgenital plate semicircular, posterior margin blunt.
Coloration  Etymology. The name of the new species is the Greek word 'nephr' (nephroid), referring to the nephroid lateral sclerites of the dorsal median lobe of the male genitalia.
Diagnosis. The new species is similar to Pseudotachycines zhengi Zhu & Shi sp. nov., but it can easily be distinguished from the latter one by the shape of the male genitalia and female subgenital plate. The dorsal sclerite of the male genitalia is semicircular and the lateral sclerite of the dorsal median lobe is nephroid. The female subgenital plate is trapezoid, with slightly concave posterior margin.
Pronotum long, anterior margin straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe with one small spine, external genicular lobe with one long spine; tibia with two inner spines and two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and two outer spines on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur with three to five inner spines on ventral surface, internal genicular lobe with one small spine, external genicular lobe unarmed; tibia with 55-57 inner spines and 55-57 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine much longer than hind basitarsus. Hind basitarsus with one dorsal apical spine.
Posterior margins of all abdominal tergites straight. Epiproct semicircular; paraproct much longer than epiproct, triangular in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia semicircular; basal area of dorsal median lobe with paired lateral sclerites, nephroid, apical area not divided; dorsal lateral lobes broad and folded, much longer than dorsal median lobe, apical area weakly sclerotized, almost as wide as basal area; ventral median lobe and ventral lateral lobes short. Subgenital plate quadrangular, posterior margin straight.
Female: Appearance is similar to male. Ovipositor longer than half the length of hind femur, curved upward, dorsal margin smooth, apical area of ventral margin denticulate. Subgenital plate trapezoid, posterior margin slightly concave.
Coloration. Dorsal surface of body light brown, with yellow spots, ventral surface yellowish brown. Eyes black.
Measurements ( Pronotum long, anterior margin straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe unarmed, external genicular lobe with one long spine; tibia with two inner spines and two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and two outer spines on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur unarmed on ventral surface, internal genicular lobe with one small spine, external genicular lobe unarmed; tibia with 47-52 inner spines and 49-50 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine nearly equal in length to hind basitarsus. Hind basitarsus with one dorsal apical spine.
All abdominal tergites without process. Epiproct nearly triangular, apex blunt; paraproct triangular in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia sagittate; basal area of dorsal median lobe with paired lateral sclerites, oval, apical area not divided; dorsal lateral lobes broad and folded, much longer than dorsal median lobe, apical area weakly sclerotized, almost as wide as basal area; ventral median Description. Male: Body medium sized in Pseudotachycines. Fastigium verticis short, with two conical tubercles, apices rounded, directing forward. Eyes ovoid, protruding forward; median ocellus oval, located between antennal sockets; lateral ocelli circular, situated on lateral margins of basal fastigium verticis. Apical segment of maxillary palpus much longer than subapical segment, apex truncated.
Pronotum long, anterior margin straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe unarmed, external genicular lobe with one long spine; tibia with two inner spines and two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and two outer spines on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur unarmed on ventral surface, internal genicular lobe with one small spine, external genicular lobe unarmed; tibia with 47-52 inner spines and 49-50 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine nearly equal in length to hind basitarsus. Hind basitarsus with one dorsal apical spine.
All abdominal tergites without process. Epiproct nearly triangular, apex blunt; paraproct triangular in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia sagittate; basal area of dorsal median lobe with paired lateral sclerites, oval, apical area not divided; dorsal lateral lobes broad and folded, much longer than dorsal median lobe, apical area weakly sclerotized, almost as wide as basal area; ventral median lobe and ventral lateral lobes short. Subgenital plate quadrangular, posterior margin arcuate.
Female: Appearance is similar to male. Ovipositor longer than half the length of hind femur, curved upward, dorsal margin smooth, apical area of ventral margin denticulate. Subgenital plate semicircular, posterior margin slightly prominent.   Etymology. The name of the new species derives from the type locality. Diagnosis. The new species is most similar to Pseudotachycines nephrus Zhu & Shi sp. nov., but it can be distinguished by the dorsal sclerite of the male genitalia semicircular, with the basal area slightly concave and the apex truncate, and the lateral sclerite of the dorsal median lobe of the male genitalia oblong.
Pronotum long, anterior margin of disc straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe with one small spine, external genicular lobe with one long spine; tibia with two inner spines and two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and two outer spines on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur with zero to four spines, internal genicular lobe with one small spine; tibia with 51 inner spines and 58 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine much longer than hind basitarsus. Hind basitarsus with one dorsal apical spine.
All abdominal tergites without process. Epiproct nearly semicircular; paraproct much longer than epiproct, oblong in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia semicircular, basal area slightly concave, apex truncate; basal area of dorsal median lobe with paired lateral sclerites, oblong, apical area not divided; dorsal lateral lobes broad and folded, much longer than dorsal median lobe, apical area weakly sclerotized, almost as wide as basal area; ventral median lobe and ventral lateral lobes short. Subgenital plate quadrangular, transverse and wide, posterior margin straight.
Female: Unknown. Diagnosis. Body is medium sized in Aemodogryllini. The male epiproct and paraproct are normally developed. The male genitalia have unpaired dorsal sclerite; the dorsal median lobe of the male genitalia has paired lateral sclerites at the basal area, the apical area does not divide; the dorsal lateral lobes of the male genitalia are little longer than the dorsal median lobe, the basal area broad, gradually narrowing to apices, the apical area sclerotized. Remarks. The species was transferred to genus Gymnaetoides according to the sclerotized dorsal lateral lobes of the male genitalia [13]. However, some characters of the species are inconsistent with Gymnaetoides: the apical area of the dorsal median lobe of the male genitalia does not divide, and the dorsal lateral lobe of the male genitalia is a little longer than the dorsal median lobe. Therefore, we transfer it into the genus Homotachycines based on morphological and molecular evidence.    Remarks. The species was moved to the genus Pseudotachycines according to the generic diagnosis [14]. However, there are some characters that differ from Pseudotachycines: the male paraproct is normally developed; the dorsal lateral lobes of the male genitalia are little longer than dorsal median lobe, with the basal area broad, gradually narrowing to apices, the apical area sclerotized, and oval lobate. Therefore, it should be transferred into Homotachycines based on morphological and molecular evidence. Remarks. The line drawing of the male genitalia of this species did not adequately represent the characters (Figure 19 in [14]). The photographs of the type specimens are shown in Figure 19. The male paraproct is normally developed. The apical area of the dorsal median lobe of the male genitalia does not divide. The dorsal lateral lobes of the Remarks. The species was moved to the genus Pseudotachycines according to the generic diagnosis [14]. However, there are some characters that differ from Pseudotachycines: the male paraproct is normally developed; the dorsal lateral lobes of the male genitalia are little longer than dorsal median lobe, with the basal area broad, gradually narrowing to apices, the apical area sclerotized, and oval lobate. Therefore, it should be transferred into Homotachycines based on morphological and molecular evidence. Remarks. The line drawing of the male genitalia of this species did not adequately represent the characters (Figure 19 in [14]). The photographs of the type specimens are shown in Figure 19. The male paraproct is normally developed. The apical area of the dorsal median lobe of the male genitalia does not divide. The dorsal lateral lobes of the male genitalia are longer than the dorsal median lobe, the basal area is broad, gradually narrowing to apices, and the apical area is sclerotized. According to these characters, this species should be moved to genus Homotachycines. The species is similar to Homotachycines ovalilobatus comb. nov. in the shape of the male genitalia, but the female subgenital plate is quite different. male genitalia are longer than the dorsal median lobe, the basal area is broad, gradually narrowing to apices, and the apical area is sclerotized. According to these characters, this species should be moved to genus Homotachycines. The species is similar to Homotachycines ovalilobatus comb. nov. in the shape of the male genitalia, but the female subgenital plate is quite different. Etymology. The name of the new species derives from the Latin word 'tri' (triple) and 'angul' (angle), referring to the triangular dorsal sclerite of the male genitalia.
Diagnosis. The new species differs from other congeneric species by the shapes of the dorsal sclerite of the male genitalia and the lateral sclerite of the dorsal median lobe of the male genitalia. The dorsal sclerite of the male genitalia is triangular, with a broadly concave apical area, and the lateral sclerite of the dorsal median lobe is quadrangular.
Pronotum long, anterior margin straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe with one small spine, external genicular lobe with one long spine; tibia with two inner spines and two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and two outer spines on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur with zero to one inner spine on ventral surface; tibia with 52-60 inner spines and 53-57 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine distinctly longer than hind basitarsus. Hind basitarsus with one dorsal apical spine.
Posterior margins of all abdominal tergites straight. Epiproct semicircular; paraproct simple, triangular in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia triangular, apical area broadly concave; basal area of dorsal median lobe with   Etymology. The name of the new species is the Latin word 'quadrat' (quadrate), referring to the dorsal sclerite of the male genitalia having a quadrate shape.
Diagnosis. The new species is similar to Homotachycines acutilobatus comb. nov., but it can be distinguished from the latter one by the apical area of the dorsal sclerite of the male genitalia slightly concave and the lateral sclerite of the dorsal median lobe of the male genitalia crescent shaped.
Pronotum long, anterior margin straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe unarmed, external genicular lobe with 1 long spine; tibia with two inner spines and two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and one outer spine on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur unarmed on ventral surface; tibia with 68-75 inner spines and 70-74 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of Etymology. The name of the new species derives from the Latin word 'tri' (triple) and 'angul' (angle), referring to the triangular dorsal sclerite of the male genitalia.
Diagnosis. The new species differs from other congeneric species by the shapes of the dorsal sclerite of the male genitalia and the lateral sclerite of the dorsal median lobe of the male genitalia. The dorsal sclerite of the male genitalia is triangular, with a broadly concave apical area, and the lateral sclerite of the dorsal median lobe is quadrangular.
Pronotum long, anterior margin straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe with one small spine, external genicular lobe with one long spine; tibia with two inner spines and two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and two outer spines on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur with zero to one inner spine on ventral surface; tibia with 52-60 inner spines and 53-57 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine distinctly longer than hind basitarsus. Hind basitarsus with one dorsal apical spine.
Posterior margins of all abdominal tergites straight. Epiproct semicircular; paraproct simple, triangular in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia triangular, apical area broadly concave; basal area of dorsal median lobe with paired lateral sclerites, quadrangular, apical area not divided; dorsal lateral lobes little longer than dorsal median lobe, basal area broad, gradually narrowing to apices, apical area sclerotized, hooked; ventral median lobe and ventral lateral lobes very short. Subgenital plate quadrangular, wide, posterior margin straight. dorsal spines and two pairs of ventral spines, intero-dorsal spine slightly shorter than hind basitarsus. Hind basitarsus with one dorsal apical spine.
Posterior margins of all abdominal tergites straight. Epiproct semicircular; paraproct simple, triangular in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia quadrate, apical area slightly concave; basal area of dorsal median lobe with paired lateral sclerites, crescent-shaped, apical area not divided; dorsal lateral lobes little longer than dorsal median lobe, basal area broad, gradually narrowing to apices, apical area sclerotized, hooked; ventral median lobe and ventral lateral lobes short. Subgenital plate trapezoidal, posterior margin straight.  Etymology. The name of the new species is the Latin word 'quadrat' (quadrate), referring to the dorsal sclerite of the male genitalia having a quadrate shape.
Diagnosis. The new species is similar to Homotachycines acutilobatus comb. nov., but it can be distinguished from the latter one by the apical area of the dorsal sclerite of the male genitalia slightly concave and the lateral sclerite of the dorsal median lobe of the male genitalia crescent shaped.
Pronotum long, anterior margin straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe unarmed, external genicular lobe with 1 long spine; tibia with two inner spines and two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and one outer spine on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur unarmed on ventral surface; tibia with 68-75 inner spines and 70-74 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine slightly shorter than hind basitarsus. Hind basitarsus with one dorsal apical spine.
Posterior margins of all abdominal tergites straight. Epiproct semicircular; paraproct simple, triangular in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia quadrate, apical area slightly concave; basal area of dorsal median lobe with paired lateral sclerites, crescent-shaped, apical area not divided; dorsal lateral lobes little longer than dorsal median lobe, basal area broad, gradually narrowing to apices, apical area sclerotized, hooked; ventral median lobe and ventral lateral lobes short. Subgenital plate trapezoidal, posterior margin straight.
Female: Appearance is similar to male. Ovipositor short, curved upward, dorsal margin smooth, apical area of ventral margin denticulate. Subgenital plate trapezoid, posterior margin straight.
Coloration. Dorsal surface of body brown, ventral surface yellowish brown. Legs with black stripes. Etymology. The name of the new species refers to the type locality. Diagnosis. The new species can be distinguished from known congeneric species by the shape of the male genitalia. The dorsal sclerite of the male genitalia is quadrangular, with the basal half rectangular and the apical half wide and slightly concave. The lateral sclerite of the dorsal median lobe of the male genitalia is oblong.
Description. Male: Body medium sized in Homotachycines. Fastigium verticis with two conical tubercles, apices rounded, directing forward. Eyes ovoid, protruding forward; median ocellus oval, located between antennal sockets; lateral ocelli circular, situated on lateral margins of basal fastigium verticis. Apical segment of maxillary palpus much longer than subapical segment, apex inflated, globular. Diagnosis. The new species is distinguished from known congeneric species by the structure of the male genitalia. The dorsal sclerite of the male genitalia is quadrate, with the basal and lateral margins slightly concave. The lateral sclerite of the dorsal median lobe of the male genitalia is fusiform.
Pronotum long, anterior margin straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe unarmed, external genicular lobe with one long spine; tibia with two inner spines and two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and two outer spines on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur with three to five inner spines on ventral surface, internal genicular lobe unarmed; tibia with 51-58 inner spines and 54-59 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs Pronotum long, anterior margin straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe with 1 small spine, external genicular lobe with one long spine; tibia with two inner spines and two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and two outer spines on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur with two inner spines on ventral surface, internal genicular lobe unarmed; tibia with 79 inner spines and 74 outer spines on dorsal surface, at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine nearly equal in length to hind basitarsus. Hind basitarsus with one dorsal apical spine.
Posterior margins of all abdominal tergites straight. Epiproct semicircular; paraproct triangular in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia quadrangular, basal half rectangular, apical half wide, slightly concave; basal area of dorsal median lobe with paired lateral sclerites, oblong, apical area not divided; dorsal lateral lobes little longer than dorsal median lobe, basal area broad, gradually narrowing to apices, apical area sclerotized, hooked; ventral median lobe and ventral lateral lobes short. transverse and wide, posterior margin straight. Female (nymph): Appearance is similar to male. Ovipositor longer than half the length of hind femur, dorsal and ventral margins smooth. Subgenital plate trapezoid, posterior margin with a concavity.
Coloration. Body yellowish brown, apex of abdomen black. Face with four longitudinal black stripes, eyes black.
Measurements (mm  Diagnosis. The new species is distinguished from known congeneric species by the structure of the male genitalia. The dorsal sclerite of the male genitalia is quadrate, with the basal and lateral margins slightly concave. The lateral sclerite of the dorsal median lobe of the male genitalia is fusiform.
Pronotum long, anterior margin straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe unarmed, external genicular lobe with one long spine; tibia with two inner spines and two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and two outer spines on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur with three to five inner spines on ventral surface, internal genicular lobe unarmed; tibia with 51-58 inner spines and 54-59 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine longer than hind basitarsus. Hind basitarsus with one dorsal apical spine.
Posterior margins of all abdominal tergites straight. Epiproct semicircular; paraproct triangular in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia quadrate, basal and lateral margins slightly concave; basal area of dorsal median lobe with paired lateral sclerites, fusiform, apical area not divided; dorsal lateral lobes little longer than dorsal median lobe, basal area broad, gradually narrowing to apices, apical area sclerotized, hooked; ventral median lobe and ventral lateral lobes short. Subgenital plate transverse and wide, posterior margin straight.
Female ( Etymology. The name of the new species is the Latin words 'concav' (concave), referring to the concave posterior margin of male subgenital plate.
Diagnosis. The new species can easily be distinguished from known congeneric species by the structure of the male genitalia. The basal two-thirds of the dorsal sclerite of the male genitalia are narrow and obtuse, the apical one-third is wide with the apex concave. The lateral sclerite of the dorsal median lobe of the male genitalia is nearly rectangular.
Pronotum long, anterior margin of disc straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur unarmed on ventral surface, internal genicular lobe with one small spine, external genicular lobe with one long spine; tibia with two inner spines and two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and one outer spine on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur with three to eleven inner spines on ventral surface, internal genicular lobe unarmed; tibia with 60 inner spines and 54 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine longer than hind basitarsus. Hind basitarsus with one dorsal apical spine.  Etymology. The name of the new species derives from the type locality, which is located in the Qinling Mountains.
Diagnosis. The new species can be distinguished from known congeneric species by the dorsal sclerite of the male genitalia trapezoidal, the basal area with a concavity, and the apical area that is widely concave; the lateral sclerite of the dorsal median lobe of the male genitalia is oval.
Description. Male: Body medium sized in Homotachycines. Fastigium verticis with two conical tubercles, apices rounded, directing forward. Eyes ovoid, protruding forward; median ocellus oval, located between antennal sockets; lateral ocelli circular, situated on lateral margins of basal fastigium verticis. Apical segment of maxillary palpus much longer than subapical segment, apex inflated, globular. All abdominal tergites without process. Epiproct semicircular; paraproct triangular in lateral view. Cercus slender, conical, apex acute. Basal two thirds of dorsal sclerite of genitalia narrow, obtuse, apical one third wide, apex concave; basal area of dorsal median lobe with paired lateral sclerites, nearly rectangular, apical area not divided; dorsal lateral lobes longer than dorsal median lobe, basal area broad, gradually narrowing to apices, apical area weakly sclerotized; ventral median lobe and ventral lateral lobes short. Subgenital plate transverse and wide, posterior margin concave.
Female (nymph): Appearance is similar to male. Ovipositor longer than half the length of hind femur, with dorsal and ventral margins smooth. Subgenital plate trapezoid, posterior margin straight.
Coloration. Body light brown, face with two longitudinal black stripes, eyes black.  Figure 20K,I and Figure 26)

Measurements (mm
dorsal spines and two pairs of ventral spines, intero-dorsal spine slightly longer than hind basitarsus. Hind basitarsus with one dorsal apical spine. All abdominal tergites without process. Epiproct semicircular; paraproct triangular in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia trapezoidal, basal area with a concavity, apical area widely concave; basal area of dorsal median lobe with paired lateral sclerites, oval, apical area not divided; dorsal lateral lobes longer than dorsal median lobe, basal area broad, gradually narrowing to apices, apical area sclerotized, apex acute; ventral median lobe and ventral lateral lobes short, cylindrical. Subgenital plate transverse and wide, posterior margin slightly concave.
Female: Unknown.   Etymology. The name of the new species derives from the type locality, which is located in the Qinling Mountains.
Diagnosis. The new species can be distinguished from known congeneric species by the dorsal sclerite of the male genitalia trapezoidal, the basal area with a concavity, and the apical area that is widely concave; the lateral sclerite of the dorsal median lobe of the male genitalia is oval.
Pronotum long, anterior margin of disc straight, posterior margin arcuate; lateral lobes longer than high, ventral margins arc shaped. Mesonotum and metanotum short, posterior margin of mesonotum arcuate, posterior margin of metanotum straight. Fore coxa with one small spine; femur with 19-25 spines on ventral surface, internal genicular lobe with one small spine, external genicular lobe with one long spine; tibia with two inner spines and two outer spines on ventral surface, apical area with one extero-dorsal spine and a pair of ventral spines, between the paired ventral spines with one small spine. Mid femur unarmed on ventral surface, internal and external genicular lobes each with one long spine; tibia with one inner spine and two outer spines on ventral surface, apical area with a pair of dorsal spines and a pair of ventral spines, between the paired ventral spines with one small spine. Hind femur with two to four inner spines on ventral surface, internal genicular lobe with one small spine; tibia with 53-57 inner spines and 55-59 outer spines on dorsal surface, with a pair of dorsal spines in subapical area, and at apex with a pair of dorsal spines and two pairs of ventral spines, intero-dorsal spine slightly longer than hind basitarsus. Hind basitarsus with one dorsal apical spine.
All abdominal tergites without process. Epiproct semicircular; paraproct triangular in lateral view. Cercus slender, conical, apex acute. Dorsal sclerite of genitalia trapezoidal, basal area with a concavity, apical area widely concave; basal area of dorsal median lobe with paired lateral sclerites, oval, apical area not divided; dorsal lateral lobes longer than dorsal median lobe, basal area broad, gradually narrowing to apices, apical area sclerotized, apex acute; ventral median lobe and ventral lateral lobes short, cylindrical. Subgenital plate transverse and wide, posterior margin slightly concave.

Phylogenetic Analyses
According to the current classification of Gymnaetoides and Pseudotachycines, the biggest difference between them is the distinctly prolonged male paraproct in Pseudotachycines [13,14]. However, the result of phylogenetic analyses shows that both Gymnaetoides and Pseudotachycines are paraphyletic, which is inconsistent with morphological classification. Therefore, we examine the type specimens of all known species and check the morphological characters of all species. In addition to the shape of the male paraproct, there are distinct differences in the shape of the male genitalia, especially in the shape of the dorsal lateral lobes and the dorsal median lobe of it. Based on the morphological characters and molecular data, we revise the genera Gymnaetoides and Pseudotachycines and a new genus Homotachycines Zhu & Shi gen. nov. is erected.
Gymnaetoides. It can easily be distinguished from Pseudotachycines and Homotachycines by the apical area of the dorsal median lobe of the male genitalia being divided into two lobes, and the dorsal lateral lobes of the male genitalia are of nearly equal length as the dorsal median lobe.
Pseudotachycines. The genus is similar to Homotachycines, but it differs from the latter one by the prolonged male paraproct and the much longer dorsal lateral lobes of the male genitalia, in which the apical area is almost as wide as the basal area.
Homotachycines. It is similar to Pseudotachycines, but it can be distinguished by the normally developed male epiproct and paraproct and the slightly longer dorsal lateral lobes of the male genitalia, in which the apical area is much narrower than the basal area.
Homotachycines is sister to Pseudotachycines, and they are together sisters to Gymnaetoides. Homotachycines and Pseudotachycines are similar in appearance to some degree. However, the dorsal lateral lobes of the male genitalia between them are distinctly different. Moreover, considering that the genus Megatachycines has developed male epiproct (quadrangular or pentagonal, much longer than the paraproct) [15], we believe that the specialized male epiproct or paraproct may be important identifying characters. Therefore, we consider Homotachycines and Pseudotachycines as two separate genera.
Last but not least, we find that the number of spines on hind femur is neither stable between different congeners nor between different individuals of the same species. Therefore, we do not think that the arrangement of spines could be a "good character" for identifying the genera, and we support Gorochov's view that the structures of the male genitalia are the more important characters [46]. As mentioned above, the classifications of genera need to be supported by more evidence and more valid characters need to be discovered.

Integrative Taxonomy
The results of the four species delimitation methods are inconsistent. The ABGD and jMOTU analyses result in 22 MOTUs, which are completely consistent with morphological species. In contrast, the bPTP and GMYC analyses generate more MOTUs. Consequently, we examine all the specimens in which results of four species delimitation methods were inconsistent. However, no distinct difference is found between different MOTUs. The more MOTUs generated by the PTP and GMYC methods might be due to the high rates of false positives of them [47][48][49][50][51].
There is also inconsistency between molecular data and morphological species. In Pseudotachycines procerus, specimens from Guizhou and Anhui form two MOTUs in all the four species delimitation methods. However, based on the morphological characters, there is only slightly difference in the lateral sclerite of the dorsal median lobe of the male genitalia, and that is not distinct enough to separate them into two independent species. Therefore, we consider specimens from Guizhou as a subspecies.
As mentioned above, 21 species and one subspecies are identified based on morphological characters and molecular evidence, of which 15 species and one subspecies are newly described: The results indicate that the molecular species delimitation methods can successfully identify most species in these genera. However, the inconsistency of morphological and genetic data or about that of different species delimitation methods suggest that species delimitation should implement multiple methods and conclusions should be drawn by integrating morphological characters and molecular data [50]. Moreover, we verify that the shapes of the dorsal and lateral sclerites of the male genitalia are suitable for the classifications of species.
To sum up, the structures of the genitalia are crucial for species diagnosis and generic identification for the three genera. In the tribe Aemodogryllini, the genitalia have sclerotized structure, while those of the tribe Diestramimini are membranous except for the genus Megadiestramima [46,52]. Therefore, Aemodogryllini is considered as a younger tribe, diverging from Diestramimini [46]. Moreover, the mating behavior of Rhaphidophoridae is unique, which may have played an important role in the evolutionary history of Orthoptera. Reproductive isolation presumable results in speciation [53][54][55][56]. The evolution of the genitalia and consequence on speciation are interesting scientific questions that can be further studied in the future.

Conclusions
Phylogenetic analyses suggest that both genera Gymnaetoides and Pseudotachycines are paraphyletic. Therefore, we revise their taxonomy based on the morphological characters and molecular data. A new genus is established, and six new taxonomic combinations are proposed. Species delimitation identifies 15 new species and one new subspecies. Moreover, we find that the shapes of the dorsal lateral lobes and the dorsal median lobe of the male genitalia are also important characters for identifying these genera, and that the shapes of the dorsal and lateral sclerites of the male genitalia are suitable for the classifications of species.
Supplementary Materials: The following supporting information can be downloaded at: https:// www.mdpi.com/article/10.3390/insects13070628/s1, Table S1: Information on the specimens included in this study and related GenBank accession numbers.