A synoptic review of the genus Thaumaspis Bolívar (Orthoptera, Tettigoniidae, Meconematinae) with the description of a new genus and four new species

Abstract Two new species of the new genus Athaumaspis gen. n., Athaumaspis minutus sp. n. and Athaumaspis tibetanus sp. n. from Vietnam and China are described. The subgenus Pseudothaumaspis of Thaumaspis is elevated to generic status and another two new species Pseudothaumaspis bispinosus sp. n. and Pseudothaumaspis furcocercus sp. n. are described, the remaining species of Thaumaspis are reviewed and keyed with the four new species.


Introduction
The genus Thaumaspis is recognized by their opisthognathous head, short wings, and completely membranous genitalia of male. The genus was proposed by Bolívar for Thaumaspis trigonurus Bolívar, 1900. Xiphidiopsis hastaticercus Tinkham, 1936and Xiphidiopsis yachowensis Tinkham, 1944 were placed into the genus by Bey-Bienko (1957). However, Gorochov (1993) thought X. hastaticercus was similar to Chandozhinskia Gorochov, 1993 without a formal assignation. Jin and Xia (1994) listed this species in Thaumaspis. We examined specimens of this species found there was no difference between Thaumaspis hastaticercus and Chandozhinskia bivittata except for length of wings and presence of some stripes, so we believe T. hastaticercus should be included in Chandozhinskia, as to whether they are the same species with different wing morphs or not needs molecular evidence. X. yachowensis was assigned to Neocyrtopsis Liu & Zhang, 2007by Wang et al. (2013. Afterwards, Thaumaspis was subdivided into three subgenera (Gorochov 1993(Gorochov , 1998) based on features of pronotum, tegmina and male genitalia: Thaumaspis s. str., Isothaumaspis Gorochov, 1993 andPseudothaumaspis Gorochov, 1998. Subsequently, Thaumaspis (Thaumaspis) henanensis Liu & Wang, 1998 (female) and Thaumaspis (Thaumaspis) bifurcata Liu, Zhou & Bi, 2010 (male) were placed in Thaumaspis both known from singletons, but now their status seems to be short of evidence and for further study the opposite sexes are required. Gorochov (1993Gorochov ( , 1998 included Thaumaspis in tribe Meconematini, but the OSF website still grouped it in Meconematinae with another 31 genera out of three tribes. Actually, those genera that were excluded from tribe Phlugidini and Phisidini should be included in Meconematini. Tirbe Phlugidini is known for their very large compound eyes, both opened protibial tympana, rounded posterior margin of pronotum, shorter tegmina than wings (except brachypterous species), ventral spines of fore femora, and short also base inflated ovipositor; Phisidini is known for their stronger ventral spines of fore and middle femora, shield protibial tympana, truncated posterior margin of pronotum, long tegmina longer than or equal to wings (except brachypterous species), and longer ovipositor (some with denticulate margins at apical half); as for Meconematini, the key features are unarmed femora, opened protibial tympana at least on one side, rounded posterior margin of pronotum, tegmina no longer than wings (except brachypterous species), and longer ovipositor with smooth margins (sometimes with an apical hook). Thus according to generic characteristics, there is no doubt for inclusion of Thaumaspis in tribe Meconematini.
Thaumaspis currently contains eight species. Here we elevate the subgenus Pseudothaumaspis to normal rank in consideration of hypognathous head and unique ventral arms of male 10 th abdominal tergite. We also describe two new species of Pseudothaumaspis, propose a new genus Athaumaspis gen. n. which includes Thaumaspis (Thaumaspis) bifurcata Liu, Zhou & Bi, 2010 for another two new species. Four new species are from China and Vietnam: Athaumaspis minutus sp. n., Athaumaspis tibetanus sp. n., Pseudothaumaspis bispinosus sp. n. and Pseudothaumaspis furcocercus sp. n.

Material and methods
The materials for this research were collected by us (from China) and came from the Bishop Museum (from Vietnam). Morphological structures were examined using a Leica MZ 125 and an OLYMUPS SZX 16 stereomicroscope, images were taken using a Motic Moticam Pro 252A digital imaging system, and drawings were produced by Adobe Photoshop from the digital images. All type specimens of new species are deposited in the SEM (Shanghai Entomological Museum, Chinese Academy of Science.) and the BPBM (Bernice Pauahi Bishop Museum, Hawaii).
In the specimen measurements, we measured length of body by distance between apex of fastigium verticis and posterior margin of tenth abdominal tergite, ovipositor by distance between base of subgenital plate and apex of ovipositor; pronotum, tegmina and posterfemora by distance between summit of base and apex. All length are shown in millimeter.

Systematics
Key to species of genus Thaumaspis Bolívar, 1900, Athaumaspis gen. n. and Pseudothaumaspis Gorochov, 1998, stat. n.  Bolívar, 1900. Description. Body small sized. Head opisthognathous. Fastigium of vertex short without sulcus dorsally, face extremely oblique, last segment of maxillary palpi longer than the preceding. Pronotum with low lateral lobes, humeral sinus absent; auditory foramina of thorax entirely exposed. Tegmina shorter than pronotum, with the stridulating organ in male, hind wing degraded. Auditory foramina of fore tibiae opened, hind tibiae with 2 pairs of apical spurs. Male 10 th abdominal tergite bearing a single process on posterior margin or absence, cerci elongate with processes, subgenital plate with short styli, genitalia entirely membranous. Female subgenital plate nearly triangular, ovipositor shorter than hind femora, ventral valve with a small apical hook.
Diagnosis. The opisthognathous head and low lateral lobes of pronotum can easily distinguished them from other genera, for now Thaumaspis is the only genus with opisthognathous head of tribe Meconematini.
Diagnosis. This new genus similar to Thaumaspis in body size and bearing posterior marginal process of abdominal tergite 10, but quite different by hypognathous head, marginal process of abdominal tergite 10 bifurcated. Description. Male. Head low in profile. Fastigium of vertex rather short, shallowly furrowed on dorsum (Fig. 24), face slightly oblique (Fig. 25), compound eyes oval and protruded outwards, last segment of maxillary palpi longer than preceding. The superior and inferior edge of pronotum nearly paralled from a lateral view, metazona slightly elevated, paranota of pronotum lower, hind margin rounded, humeral sinus absent; auditory foramina of thorax entirely exposed. Tegmina shorter than pronotum, hind margin obliquely truncated, hind wings reduced. Fore tibiae armed spines of type 4, 4 (1, 1) on either margin of ventral surface, hind tibiae with 20-23 dorsal teeth each margin above and 2 pairs of apical spurs. Posterior margin of abdominal tergite 10 with middle process, distinctly branched (Fig. 26). Epiproct reduced. Cerci elongate, rather simple, incurved in its apical third, dorsal surface with weak keels (Fig. 27). Subgenital plate narrowed basally, broad in apical half, hind margin roundly emarginate with rather short styli (Fig. 28).
Coloration. Body yellowish (maybe greenish in life), eyes blackish brown, antennae with inconspicuous darkish rings, fore and hind margins of pronotum either with blackish brown marking rounded yellow rim (Fig. 24), lateral lobe with 4 yellow markings.
Diagnosis. This new species distinguishes from other species of the genus in body smaller, pronotum with blackish brown and yellow markings, female subgential plate with rounded posterior edge.
Etymology. The specific epithet referrers body form of this species, from Latin minūtus. The gender of the epithet is masculine.
Distribution. Vietnam. Description. Male. Head high in profile. Fastigium of vertex short, dorsum shallowly furrowed in middle, face slightly oblique (Fig. 31), but higher, compound eyes subovoid and moderately protruded, last segment of maxillary palpi slightly longer than preceding. Protonum nearly triangular in profile, metazona little elevated, paranota higher, hind margin obliquely truncated without humeral sinus; auditory foramina of thorax entirely exposed. Tegmina shorter than pronotum by one third, posterior edge truncated; hind wings deduced. Fore tibiae spines armed 4, 4 (1, 1), hind tibiae with 19-20 dorsal teeth either margin above and 2 pairs of apical spurs. 10 th abdominal tergite bearing an extended process at the middle of hind margin, bending vertically downwards and invisible dorsally (Fig. 32), apex distinctly branched (Fig. 33). Epiproct reduced. Cerci longer, inner surface of base occurs a lobe, incurved in one third, apex moderately expanding. Subgenital plate longer than width, apical two fifth narrowing towards tip, little convex at median hind margin, styli short (Fig. 34).
Diagnosis. This species looks different from type species in general, distinguishes mainly by higher head and pronotum in profile and inner lobe of cerci; but shearing branched process of male 10th abdominal tergite and simple but slender cerci.
Etymology. The specific epithet is Latinized name of district Tibet where this species distributed. The gender of the epithet is masculine.
Distribution. China (Xizang).  Diagnosis. Posterior margin of male 10 th abdominal tergite with a small branched process, male cercus stout and bifurcated, superior ramus clubbed, inferior ramus with an inner triangular lobe at proximal part.

Athaumaspis bifurcatus
Coloration. Body yellowish green. Eyes and spines of hind tibiae blackish, genicular lobes of all femora each with a blackish spot.

36
Discussion. The cerci of this species are stouter and quite different from previous 2 species of this genus which makes it easy to tell them apart, meanwhile the species meets the diagnosis of Athaumaspis in hypognathous head and bifurcated posterior marginal process of abdominal tergite 10. According to the features of male here we treat this species as an Athaumaspis. The specific epithet of this species originally was feminine, primarily based on Gorochov (1993), but according to type species of Thaumaspis the genus is masculine and 'bifurcata' should be 'bifurcatus', The same apply for Athaumaspis and Pseudothaumaspis.

Pseudothaumaspis gialaiensis
Coloration. Body yellowish green, almost unicolor, antennae with brown rings, lower part of the pronotum lateral lobe with brown edge; apex of tibiae, tarsus and spine of tibiae darkened.  Gorochov, 1998 (after Gorochov)  Description. Male. Fastigium of vertex short, without dorsal groove, face slightly oblique (Fig. 42), compound eyes oval and protruded forwards and outwards, last segment of maxillary palpi slightly longer than preceding. The superior edge and inferior edge of pronotum paralleled and almost straight, paranota lower, front margin little sinuate, hind margin straight without humeral sinus and obliquely truncated; auditory foramina of thorax small and entirely exposed. Tegmina almost equal to pronotum, apex rounded; hind wings degenerate. Fore tibiae armed ventral spines of type 4, 4 (1, 1), lower lobe of the hind knee bearing a spine, hind tibiae with 28-31 dorsal teeth each margin above and 2 pairs of apical spurs. Hind margin of 10 th abdominal tergite little sinuate (Fig. 43), lower part becoming a pair of elongate branches (Figs 44-45); cerci robust, generally conical and apex blunt, but each with 2 long inner processes: prior one downward, posterior one upward and little branched at apex. Subgenital plate damaged.
Female unknown.
Discussion. This new species is similar to Pseudothaumaspis gialaiensis Gorochov, 1998, but differs mainly in the appearance of the male cerci. The single specimen of this species is in bad condition, especially the abdomen. Fortunately, the unique ventral arms of the 10 th abdominal segment and the cerci are intact.
Etymology. The specific epithet refers to the character of male cerci which bearing 2 spinous processes, compose by prefix 'bi-' which means double and 'spinosus' which means spiny.
Distribution. Vietnam.   (Fig. 48), compound eyes subglobular, last segment of  maxillary palpi slightly longer than preceding. Pronotum saddle shaped in lateral view, paranota with concave dorsal margins, ventral margin rounded, humeral sinus absent, transverse sulcus distinct especially posterior one, metazona short, rather pointed at the posterior tip; auditory foramina of thorax small and exposed. Tegmina shoter than pronotum by one third, apex truncate; hind wings degenerate. Legs very long and rather thin (Fig.  46), fore tibiae with ventral spines armed 4, 4 (1, 1), lower lobe of the hind knee without spine, hind tibiae with 21-29 dorsal teeth either margin above and 2 pairs of apical spurs. Posterior median edge of 10 th abdominal tergite elongate with a deep apical incision forming 2 small lobes (Fig. 49), lower area bearing a pair of ventral arms, not elongate and apex oblate (Fig. 52). Slim cerci branched at a half, incurved, lower branch longer. Subgenital base broad, apical one third narrow and up curved, styli very long (Figs 50-52).
Female. Body form similar to that of male. Fastigium of vertex little longer and more slender, a shallow furrow on the dorsum, last segment of maxillary palpi much longer than preceding. Paranota of pronotum subacute at inferior margin, transverse sulcus distinct as in male, but metazona even shorter. Tegmina short as in male, pointed at apex, inferior margin obliquely truncated, veins conspicuous. Posterior margin of 9 th abdominal tergite straight, cercus slender, fusiform, apex thin and acute (Fig. 54); subgenital plate downward swell, trilobed at hind margin, mesolobe prominently convex. Ovipositor short, base upcurved, ventral valve without apical hook.
Coloration. In life of male. Body lightish green, emerald green and lightish yellow variegated. Flagella pale brown with darkish rings, scape and pedicel consistent with body color. Compound eyes vivid yellow. Both lateral rims of pronotum emerald green, but posterior edge vivid yellow, dorsum with green longitudinal stripes and patches. Each abdomen tergite with a pair of bright yellow oval patches and posterior edge darkish green. Hind tibiae, Tarsi and cerci terminal pale brown.
Dry specimen. Body brownish, antennae with inconspicuous darks rings, forma and tibia darkened around the knee joint. Male unicolor; female abdomen largely blackish brown, ventral surface totally black including subgenital plate, abdomen tergites each compact with a pair of large pale patches dorsally, base of ovipositor darkened.
Etymology. The specific epithet from Latin 'forca' + 'cercus', corresponding the feature of male bifurcate cerci. The gender of the epithet is masculine.
Discussion. Bearing those unique arms, clearly it is a Pseudothaumaspis, but tegmina terminal, lower lobe of hind knee, small lobes of posterior edge of last abdominal tergite are quite different from previous species.