Notes on the genus Theopropus Saussure (Mantodea, Hymenopodidae) from China, with description of a new species from the Himalayas

Abstract The genus Theopropus Saussure, 1898 is distributed with five species in SE Asia, three of which were recorded in South China: T. elegans (Westwood), T. cattulus (Westwood) and T. sinecus Yang. After examining numerous specimens of Theopropus, we attempt to resolve some taxonomic confusion about Theopropus occurring in China. Those Chinese specimens that were inaccurately identified as T. cattulus represent a new subspecies of T. sinecus Yang: T. s. qiongae Wu & Liu, ssp. nov.. A new species, T. xishiae Wu & Liu, sp. nov., is described from the rainforests of the southern slopes of the Himalayas. The records of T. elegans in China are also clarified. Biological characteristics of the species and subspecies, necessary illustrations, and ecological images are provided. The distribution of the known Chinese Theopropus species is discussed and mapped.

Theopropus is widely distributed in southern China, the Indochinese Peninsula, and the Malay Archipelago. In China, T. elegans was first mentioned to be distributed in Yunnan by Tinkham (1937), but without specimen records. The report of T. elegans from Yunnan in Tinkham (1937) was questioned by Wang (1993) and Yang and Wang (1999) because no specimens were examined. Afterwards, a male specimen of T. elegans was reported from Wuyishan Mountain in Fujian Province by Wang (1993). Subsequently, the new species T. sinecus was described by Yang (1999) based on a female (holotype) and a male (paratype) specimen, which were collected from Jinxiu in Guangxi Province. Additionally, T. cattulus Westwood was reported to be distributed in Hainan Island by Zhu et al. (2012), who also noted that T. sinecus was a synonym of T. elegans, but without standard taxonomic treatment. As a consequence, there is considerable taxonomic confusion concerning the common and attractive mantis genus Theopropus in China.
In this research, we examined numerous specimens, which were collected in China and neighboring countries, aiming to illustrate the taxonomic situation of the genus Theopropus in China. We clarified the validity of Theopropus sinecus Yang, the distributions of T. elegans and T. cattulus, redescribed the known Chinese species and describe a new taxon.

Materials and methods
Classification system follows Schwarz and Roy (2019). Descriptive terminology of adult morphology and the male genitalia follows Brannoch et al. (2017) and Schwarz and Roy (2019). Specimens were collected during the daytime through careful observation or by light trap (male). Genitalia were dissected in 10% KOH solution, cleared with pure water, and finally stored in 70% ethanol in Eppendorf tubes for further research. Pictures were taken with a Nikon digital camera.
Type species. Blepharis elegans Westwood, 1832 Diagnosis. Medium to large-sized Hymenopodidae, with mottled body coloration. Male and female distinctly differing by body size, male body smaller, often shorter than half body length of females. Head (Fig. 5): Triangular. Compound eyes oval, convex, uprising beyond vertex. Vertex with a robust vertical process, coniform. Lower frons wider than high. Antennae filiform, shorter than body length; antennae thick and long in males, thin and short in females.
Meso-and metathoracic legs: Long, robust; subapical part of the femur with a posteroventral lobe (Fig. 9D-F). Base half of tibia swollen.
Wings: Forewings opaque, narrow, long in males, wide, fusiform in females; a white spot lying subbasally in the discoidal area; a white band with black borders on both lateral margins lying in middle of the discoidal area; anal area long, narrow. Hindwings broad, shorter than forewings; transparent or with opaque areas in males, subopaque in females.
Abdomen: Narrow, long in male, wide in female. Cerci short, hairy. Male subgenital plate short, wide, with small styli.
External genitalia (Fig. 10A-F): Male genitalia simple, similar among congeners. Secondary distal process reduced. Ootheca (Fig. 4C): Very elongated, flat, narrowing at both ends. Discussion. The characteristics for the head, pronotum, and the range of the ratio of the pronotum length to supracoxal dilatation width are relatively stable in the species; these characteristics can be used to identify species. The male genitalia of Theopropus lack sclerotized projections and show little differences between species.
Distribution. The genus Theopropus is distributed in the tropical areas of southern Asia. In China, Theopropus species are widely distributed in South and Southwest China (Fig. 12).

Theopropus sinecus Yang, 1999
Theopropus sinecus Yang, 1999: 28;T. elegans Zhu et al. 2012: 52-55. Comments. Theopropus sinecus Yang is widely distributed in South China. Its types were collected from Guangxi Province. Specimens from the mainland and Hainan Island have similar body features and were mated to produce fertile offspring in our own breeding facilities, but they clearly differ by body color and spots' characteristics in the forewings. Therefore, we consider specimens from Hainan Island as a new subspecies for T. sinecus Yang. Redescription. Male. Large-sized compared with congeners, body length reaching half that of females.  Head: Compound eyes oval, anteriorly protruding, with a very weak conical elongation at tip (Fig. 5A). Vertical process distinct, coniform, lower than the apex of compound eyes (Fig. 5A). Lower frons with arched superior margins and raised lateral margins. Antennae filiform, shorter than body length.
Female. Similar to male, but body more robust, size larger than that of male. Vertical process distinct, conical, extending beyond apex of eyes (Fig. 5D). Pronotum wide, lateral pronotal expansion very wide, ratio of pronotum length to supracoxal dilatation width about 1.37-1.40; black band on each lateral margin of metazone traversed by a white band (Fig. 7A). Forewings wide, fusiform, extending beyond abdominal apex. Hindwings broad, opaque. Abodomen broad, nearly rounded, 4 th -6 th abdominal tergite inconspicuously slightly expanded laterad.
Coloration. Overall green, with white spots and bands. Antennae black. Lateral margin of metazone band black. The two horizontal ventral bands on prothoracic femora and tibia black in male (Fig. 8A), absent on femora for females (Fig. 8D). A black ventral spots near prothoracic tibial spur (Figs 6A, 7A). Forewings green, costal area white, discoidal area green; the large subbasal spot of discoidal area white, with black hind edge; wide band in middle of discoidal area white, with black lateral margins; anal area orange in males and white in females. Hindwings orange with red-brownish veins in males, whitish to slightly yellowish with hyaline margin in females. Abdomen yellowish white, plaques on lateral margins of 2 th -6 th abdominal tergite green (Fig. 9B).
Note. When examining numerous specimens from Wuyishan, Fujian Province (listed above), we found that they are the same as the types of Theopropus sinecus. Their body is larger than in T. elegans from Malaysia; the compound eyes possess a conical elongation at the tip in males; the black band on each lateral margin of the pronotum is continuous in males, but disconnected in females. In T. elegans, the compound eyes do not have a conical elongation in males, and the black band on the lateral margin of the pronotum is contiguous in both sexes. Also, female hindwings are orange with smoky margins in T. elegans. The specimens of Theopropus from Wuyishan should therefore be identified as T. sinecus sinecus. We think that the record of T. elegans (in Wuyishan, Fujian Province) might have been mistaken by Wang (1993) and Wang and Yang (1999). Yang (1999) wrote the specimen information in the Chinese description as "Guangxi-Dayaoshan Mt., 1981-VIII, Qijing You leg.", however, the specimens of Theopropus with the same collection information could not be found among Yang's research specimens. Only one female labeled "Guangxi, Jinxiu; 18-XI-1981" and one male labeled "Guangxi, Jinxiu; 21-IX-1981" could be found. We re-measured these specimens and obtained the following measurements: length of body (head to wings) about 29.3 in male and 45.2 in female, length of forewings about 21.6 in male and 29.2 in female (impaired), in original description, length of body (head to wings) 31 in male and 44 in female, length of forewings about 22 in male and 29 in female. In addition, as the illustration (hand-painted) of the original description is similar to the posture of the female specimen, we confirm that these two specimens are the types of T. sinecus Yang (Figs 4A, B, 10F), except the following characteristics: smaller; on surface of forewing, the white subbasal spot of discoidal area small, even disappearing in some specimens; white band at the middle of discoidal area narrow, the two black parallel arc-shaped lateral borders of the white band converging at their distal ends in rare instances; hindwing base reddish brown, with red-brownish veins.
Female. Similar to T. s. sinecus but smaller. The forewing is similar to male, white spot in base of discoidal area small or indistinct, white band in middle of discoidal area narrow; hindwings opaque, maize-yellow, with hyaline margin.
Discussion. Zhu et al. (2012) identified the specimens from Hainan Island as Theopropus cattulus (Westwood, 1889). After checking the pictures of the types of T. cattulus Westwood, 1889 in "The Mantodea Image Database" https://specimens. mantodearesearch.com/default/zoom/835, we found that there is only a small black spot on each lateral margin of the metazone in T. cattulus, whereas, in those specimens from Hainan Island, a black band extends backwards on each lateral margin of metazone. Also, Theopropus cattulus is endemic on Java, Indonesia. Thus, we think the identification of these specimens from Hainan Island in Zhu et al. (2012) is wrong. We establish a new subspecies of T. sinecus for these specimens from Hainan Island.
Head: Compound eyes oval, anteriorly protruding, with rounded top. Vertical process conical, extending about as high as the imaginary line between the apexes of the eyes; lower frons narrow, with arched dorsal margin and raised lateral margins (Fig. 5B). Antennae filiform, shorter than body length. Pronotum (Fig. 6B): Wide. Lateral pronotal expansion very wide; lateral margins bearing small, sparsely arranged spines. Black band on each lateral margin of metazone disconnected in middle. Ratio of pronotum length to supracoxal dilatation width about 1.39-14.2.    Forewings: Long, narrow, opaque. Discoidal area possessing a large subbasal white spot with blurry black edges; the two black lateral borders of the wide white band in middle of the discoidal area arched, the anterior margin of the frontal one directing proximad, of the caudal one directing distad (Fig. 13D). Anal area long, narrow, hyaline.
Female. Large-sized, robust. Body size largest among known Theopropus species. Head (Fig. 5E): Similar to male, but vertex extending beyond apex of eyes. Pronotum (Fig. 7B): Wide. Lateral pronotal expansion very wide. Lateral margins bearing small, sparsely arranged spines. Black band on each lateral margin of metazone disconnected in middle. Ratio of pronotum length to supracoxal dilatation width about 1.08.
Forewings (Fig. 3B): Wide, fusiform, opaque. The large white subbasal spot of the discoidal area with black edges; frontal one of the two black lateral margins of the wide white band in the middle of discoidal area arc-shaped, pointing proximad, and hind one approximately straight. Anal area long, narrow, hyaline.
Coloration. Yellowish green, with white spots and bands. Antennae black. Band on each lateral margin of metazone black. Two horizontal ventral bands on prothoracic femora and tibia black in males (Fig. 8B), which is absent on femora in females (Fig. 8E). Forewings yellowish green, costal area white; the large spot in base of discoidal area white, with black hind edge; the wide band in middle of discoidal area white, with black lateral margins; anal area orange in males and white in females. Hindwings hyaline, with red-brownish veins in males, ivory in females. Abdomen yellowish white; lateral margins of 3 th -5 th abdomen tergite with green plaques, and 6 th -7 th mostly white in females.
Distribution. China: Tibet, Medog. Expected to also occur in N India. Etymology. The new species was named after Xi Shi who was born in The Spring-Autumn Period, the top of the four beautiful women in ancient China, the beauty representative in Chinese culture. Comments. Male. Compound eyes oval, anteriorly protruding. Prolongation bifid vertex conical, not reaching imaginary line extending between the apexes of the eyes (Fig. 5C). Lateral pronotal expansion wide, ratio of pronotum length to supracoxal dilatation width about 1.51-1.53. Lateral margins of pronotum bearing inconspicuous teeth. Black band on each lateral margin of metazone continuous (Fig. 6C). Anterior coxa bearing 8-10 dorsal spines, femora with 17 anteroventral spines, tibia with 17-18 anteroventral and 21 posteroventral spines; tibia with two black horizontal bands on ventral side (Fig. 8C, F). Forewings green; the black lateral borders of the white band in middle of discoidal area wide, blurry. Hindwings orangish red, transparent.

Differential diagnosis.
Compared with the other two species of Theopropus in China, this species is smaller in body size, the prolongation on the vertex is small in the female, and the difference in body size between the sexes is more pronounced. The dorsal spines on anterior coxae are larger and longer than those of the other two species. The anterior tibia does not have a black spot near the spur. The male characteristics are also close to that of T. cattulus (Westwood, 1889) (type locality in Java, Indonesia) but the markings of the pronotum and forewings are different. In addition, in this species, the male hindwings do not have the opaque area which is present in the male specimens of T. elegans from the Malay Peninsula. These specimens may represent another new species, and further research on this species is needed.
Distribution. China: Yunnan; Thailand. Biological characteristics. Theopropus species often live among flowers. In Huaping of Guangxi Province (southwestern of China), T. sinecus sinecus often appears among the inflorescences of Valerianaceae plants, the mottling pattern of the mantis allows them to blend in such an environment (Fig. 13A, B). The males have phototaxis during night time.   In China, the genus Theopropus ranges northwestwards to Medog, Tibet, and eastwards to the central Fujian Province. Theopropus species often inhabit mediumelevation forests. In Guangxi Province, T. sinecus sinecus is distributed at an altitude of about 800-1400 m. Theopropus sinecus qiongae ssp. nov. was collected from 800-1000 m in Hainan Island. Theopropus species in Yunnan Province were collected from 600-1100 m. Theopropus xishiae sp. nov. from Tibet were collected from 900-1400 m. Theopropus sinecus sinecus overwinters as eggs or nymphs in Guangxi, Fujian, and Guangdong Provinces. In the mountains of these areas, it snows in winter, and the lowest temperature about -5~-10 °C. Nymphs of T. sinecus sinecus hide in the deciduous layers during winter, and begin to grow about April of the following year; adults can be seen from July to November. In Hainan Island, no clear seasonality patterns can be observed in T. sinecus qiongae ssp. nov., for which adults and nymphs can be found in each season. The same situation is found in southern Yunnan. The situation for T. xishiae sp. nov. in Medog of Tibet is unclear, but adults of T. xishiae can be seen from July to October.
In rare instances, females of Theopropus sinecus collected from Guangdong and Hainan have been discovered to be parasitized by horsehair worms.

Discussion
Hainan Island is an isolated island in southeastern China. The Qiongzhou Strait between Hainan Island and the mainland is a geographical barrier for the separation and differentiation among closely-related species which separately live on either side of the barrier, as is the case postulated for T. s. sinecus and T. s. qiongae ssp. nov.. The ancestor of T. xishiae sp. nov. in the Himalayas is speculated to come from the Assam Region, and numerous mountains have become obstacles which prevented them from spreading northwards. However, in southern Yunnan, the distribution boundaries of T. s. sinecus and T. sp. are not yet clear, the two species seem to be separated in the Honghe area; T. s. sinecus lives in the eastern part, and T. sp. lives in the western part.
The structure of the male genitalia of Theopropus is rather simple compared to other species within the order Mantodea. Nonetheless, the shape of the ventral phallomere can be used to distinguish the three species in China: the ventral phallomere is wide and secondary distal process (spd) indistinct in T. sinecus; it is wide and with inconspicuous secondary distal process in T. xishiae sp. nov.; and it is rhomboidal in T. sp.

Conclusion
After examining numerous specimens, which were collected in China and neighboring countries, we reached the following conclusion. Theopropus sinecus Yang is valid. Those specimens that were collected from Hainan Island and identified as T. cattulus Westwood by Zhu et al. (2012) belong to a new subspecies for T. sinecus, i.e., T. sinecus qiongae ssp. nov.. The male specimen, which was identified as T. elegans in Wuyishan, Fujian Province by Wang (1993) should also be identified as T. sinecus. A unique new species, T. xishiae sp. nov. was discovered in the Himalayas (Tibet in China). In addition, numerous specimens of Theopropus from southern Yunnan are temporarily assigned to an unidentified species not identical with the previous two, and also not identical with T. elegans. The species T. elegans is not distributed in China. also express our thanks to Dr. Roberto Battiston (Italy) and Christian J. Schwarz (Germany) for their careful reviews and valuable comments. The study was funded by the National Natural Science Foundation of China (No. 31572308, 31750002).