Two new species and a new combination of Allacta (Blattodea, Ectobiidae, Pseudophyllodromiinae) from China, with notes on their behavior in nature

Abstract Two new species, Allactabrunasp. n. and Allactaalbasp. n., from China are described and illustrated. Allactahainanensis (Liu et al., 2017), comb. n. is proposed and re-described; figures including genitalia are provided. A key is provided to all species from China based on males. Notes on the bionomics of this genus in China are provided.


Introduction
The genus Allacta is widely distributed from southwestern China to northeastern Australia. It consists of 41 known species worldwide, five of which were known from China before this study. Wang et al. (2014) described the species Allacta xizangensis, re-described four other known species from China, and distinguished Allacta from related genera. Liu et al. (2017) described a species Temnopteryx hainanensis from Hainan, China, based on morphological characters (without providing male genital characters), and which has reduced tegmina and wings. However, Princis (1963) proposed Temnopteryx is endemic to South Africa. Princis also established the generic diagnosis as follows: tegmina absent or shortened, hind wing reduced to lateral lobes or absent; tarsal claws symmetrical and unspecialized; and abdominal terga unspecialized. Fortunately, we obtained specimens of T. hainanensis from Hainan Province as well as a photograph of the holotype. We also collected specimens from Zhejiang and Hainan, among which two new species of Allacta were discovered, and we use this opportunity to describe them together. With an increase in field surveys, it has been possible to better observe the behavior in nature of Allacta species and the members of this genus are commonly found to be tree climbers in the forests.

Materials and methods
Terminology used mainly follows Roth (2003). Genitalia terms are according to McKittrick (1964). Venation terms follow Li et al. (2018). Vein abbreviations in this article are as follows: Measurements are based on specimens examined. The genital segments of the examined specimens were soaked in 10% NaOH and rinsed with distilled water, then stored in glycerin for observation. Genitalia were observed in glycerin using a MOTIC K400 stereomicroscope and genital photographs were taken using an LAS V4.9. Specimen photographs were taken using a Canon 50D plus a Canon EF 100mm f/2.8L IS USM MACRO lens and photos were stacked using Helicon Focus software. All photos were modified in Adobe Photoshop CS6. The materials examined are deposited in the Institute of Entomology, Southwest University, Chongqing (SWU) and the Shanghai Entomological Museum, the Chinese Academy of Sciences, Shanghai (SHEM). Diagnosis. This species can be easily distinguished from its congeners by the brownish body that lacks any markings.
Vertex with interocular space about half the distance between antennal sockets (Fig. 3). The third, fourth and fifth maxillary palpi of approximately same length (Fig. 5). Pronotum nearly trapezoidal, broader than long, maximum width behind the middle, the front and hind margins nearly straight, and the postero-lateral angle blunt and round (Fig. 4). Tegmina and wings fully developed, both extending beyond the end of abdomen. Tegmina with M and CuA longitudinal (Fig. 7). Hind wings with M straight and simple; CuA slightly curved with 4 branches (Fig. 8). Anteroventral margin of front femur type B 2 (Fig. 6). Pulvilli only present on the fourth tarsomere. Tarsal claws symmetrical and unspecialized, arolium present. The first tarsus of the front legs about equal in length to the sum of the other four, but obviously longer in the middle and hind legs.
Female similar to the male. The coloration of living individuals dark brown, dried specimens much lighter, especially in the legs. Margins of pronotum and tegmina subtransparent in the specimens.
Male abdomen and genitalia. Abdominal terga unspecialized. Supra-anal plate nearly triangular, and the hind margin obviously concave. Paraprocts simple (Fig. 9). Subgenital plate nearly symmetrical, the middle part of the hind margin has a V-shaped concavity; styli similar, short cylindrical, with small spines, and interstylar margin with two lobes (Fig. 11). Left phallomere complex, inverted Y-shaped (Fig. 10). Median phallomere stem club-like, apex shoe-shaped, base with a small spine, the accessory structure curved, apex brush-shaped (Fig. 11). Hook on the right side (Fig. 12).
Remarks. It can be distinguished from the other species in the group by the tegmina that without any light-colored bar in the anal field.
Etymology. From the Latin word brunus referring to the body color being brown without any special markings.
The distance between compound eyes narrower than that of antennae, ocelli border not obvious (Fig. 15). Curved macula under the antennal sockets. Third, fourth and the fifth maxillary palpi approximately same length (Fig. 17). Pronotum nearly triangular, width greater than length, the front and hind margins nearly straight, the posterolateral angle blunt and round, and disc with trapezoidal symmetrical white maculae (Fig. 16). Tegmina and hind wings fully developed, both extending beyond the end of abdomen (Fig. 13). Tegmina with M and CuA longitudinal, CuA with four branches (Fig. 19). Hind wings with M straight and simple; CuA with five complete branches (Fig. 20). Anteroventral margin of front femur type B 3 (Fig. 18). Pulvilli present only on the fourth tarsomere; tarsal claws symmetrical and unspecialized, arolia present.
Remarks. It can be distinguished from other species in this group by the lightcolored pronotum.
Etymology. The meaning of the Latin word alba is white, referring to the white maculae on the pronotum.
Distribution. China (Zhejiang). (  Diagnosis. This species can be easily distinguished from all other congeners by the much reduced tegmina and wings, which only reaching the third tergum. The coloration pattern (body light yellowish brown, with large brown areas dorsally) is also unusual in this genus (see Figs 25 and 45).
Description. Male. Body dark brown with yellowish (Fig. 25). Frons yellowish brown, vertex with dark brown bands. Antenna brown. The fifth of maxillary palpomere brown (Fig. 26). Pronotum yellowish brown, lateral and hind border translucent, and disc with two dark brown stripes and an irregular pale yellowish brown macula. Tegmina brown, the inner border dark brown, lateral border pale yellowish brown (Fig. 25). Legs pale yellowish brown. Abdominal terga brown, lateral margin with blackish brown spots. Cerci yellowish brown, base brown (Fig. 26).
Vertex with interocular space obviously narrower than the distance between antennae sockets (Fig. 27). Third and fourth maxillary palpomeres approximately same length, and both significantly longer than the fifth. Pronotum nearly triangular, the front margin blunt and round; the hind margin nearly flat (Fig. 28). Tegmina and hind wings reduced; tegmina nearly quadrilateral, veins inconspicuous, only reaching the third tergum of the abdomen (Fig. 30); hind wings short and narrow, about half the length of the tegmina, and veins simple with two longitudinal veins (Fig. 31). Anteroventral margin of front femur type B 3 (Fig. 29). Hind legs first tarsus approximately same length to the sum of other four tarsomeres. Pulvilli present only on the fourth tarsomere, tarsal claws symmetrical and unspecialized, arolia present (Figs 25,26).
Female similar to the male. Male abdomen and genitalia. Abdominal terga unspecialized. Supra-anal plate nearly triangular, symmetrical, the middle of hind margin with incisions. Paraproct plates simple, similar, sheet-like, apex with scattered bristles (Fig. 32). Subgenital plate symmetrical, lateral margin curved and blunt; styli nearly cylindrical, arising in two concavities of hind margin; the right stylus slightly longer than the left; and the hind margin with W-shaped concave (Fig. 34). Left phallomere complex (Fig. 33). Median phallomere stem slender, bending near apex, apex sharp, base forked, median phallomere subsidiary sclerite C-shaped clavate, apex gradually sharper (Fig. 35). The hook-shaped phallomere on the right of subgenital plate, and the hook short (Fig. 36).
Remarks. Liu et al. (2017) placed this species in Temnopteryx and stated it resembles T. dimidiatipes Bolivar, 1890 from the Philippines. However, Princis (1957) erected Lobopterella and treated T. dimidiatipes as the type species. Thus Lobopterella dimidiatipes belongs to subfamily Blattellinae and does not display the characteristic of Pseudophyllodromiinae, viz. the hook-shaped phallomere on the right side (Roth 1988).
Through field efforts, we obtained several specimens with reduced tegmina and wings from Hainan Island. After comparing them with the type specimen (deposited in SHEM), we confirmed that our specimens are Temnopteryx hainanensis Liu et al., 2017. After dissecting the male genitalia, we found the hook-shaped phallomere of Temnopteryx hainanensis is on the right, and the pulvilli is present only on the fourth tarsomere, thus, Temnopteryx hainanensis should be placed in the pseudophyllodromiine genus Allacta as Allacta hainanensis (Liu et al., 2017) comb. n.
This species is placed in the hamifera species group by having two lobes which forming a keel-like ridge, and in having the dark portion of the pronotum reduced to two longitudinal bands.

Discussion
Previous studies indicated that Allacta species inhabit on tree trunks (Rentz 2014;Wang et al. 2014). Through our field investigations from 2014 to 2017, we found that most Chinese Allacta species are more likely to inhabit tree trunk surfaces or under the bark. Sometimes, individuals are also observed crawling on the leaves (Fig. 45) or ground (Fig. 44). A. bimaculata, A. ornata, A. robusta, A. transversa, A. bruna sp. n., and A. xizangensis are observed resting on tree trunks. Their bodies are maculated (except for A. bruna sp. n.), and some species even have reticulated mottling on the tegmina (e.g., A. bimaculata, A. ornata, A. transversa), which provide an excellent disguise on a tree trunk surface that is covered with lichens and mosses (Figs 37-43). In the daytime, Allacta individuals may hide under the bark (Fig. 37), while during the night, they will come out and crawl on the trunk. When approaching them, they would not act dead or fall down to the ground, but instead hide from the collectors. They can move transversely somewhat like crab and very rapidly slip away to the opposite side of the trunk to escape from being captured. While when the collectors continue tracking them, they will move around to the other side of the trunk, or go to a higher level on the trunk. Characters such as the presence of pulvilli only on the fourth tarsomere and spines on the other 3 tarsomeres help Allacta species efficiently move about on the trunk. They can conduct such fast transverse actions probably because they only walk or climb "on tiptoe" (Wang et al. 2014).