Three new species of Pseudophanias Raffray from Japan and Taiwan Island, and synonymy of Chandleriella Hlaváč with Pseudophanias (Coleoptera, Staphylinidae, Pselaphinae)

Abstract The genus Pseudophanias Raffray, 1890 is discovered in Japan and Taiwan Island for the first time, with three new species: P. yaimensis Inoue, Nomura & Yin, sp. nov., P. nakanoi Inoue, Nomura & Yin, sp. nov., and P. excavatus Inoue, Nomura & Yin, sp. nov. It is the fifth tmesiphorine genus known from Japan and the first from Taiwan. The genus Chandleriella Hlaváč, 2000 is placed as a junior synonym of Pseudophanias, resulting in the following new combinations: P. termitophilus (Bryant, 1915), comb. nov., and P. yunnanicus (Yin, 2019), comb. nov. A list of world species, and a key to East and South Asian representatives of Pseudophanias is provided.


Introduction
The tribe Tmesiphorini Jeannel currently contains 30 extant genera worldwide (Yin et al. 2013a), among them the primarily Oriental genus Pseudophanias Raffray, 1890 was established for P. malaianus Raffray from Malaysia (Raffray 1890a, b). Pseudophanias was initially placed in the tribe Tyrini (Raffray , 1908, then in Phalepsini (Jeannel 1949;Newton and Chandler 1989), and was most recently moved to Tmesiphorini (Chandler 2001). Currently, ten epigean species are known from the tropical areas in Southeast Asia (Raffray 1890a(Raffray , b, 1895, and one cavernicolous species was described from central Nepal . Hlaváč (2000) erected a new genus Chandleriella Hlaváč for Lasinus termitophilus Bryant from Borneo. He correctly recognized Chandleriella as a member of Tmesiphorini based on the presence of semi-circular sulci that enclose the antennal bases, but did not compare it to any members of the tribe. A second species of the genus, C. yunnanica Yin, was later described from southwestern China , with two individuals of the type series associated with Ectomomyrmex ants.
Specimens of the genus Chandleriella have well-developed posterior tarsal claw and an overall elongate body form, whereas those of Pseudophanias have a strongly reduced posterior tarsal claw and a stouter body. However, after examining the type species of Pseudophanias (by the second author), and a vast collection of undescribed Pseudophanias-Chandleriella-like species, transitional states in both claw morphology and habitus were found. Nomura and Idris (2005) already discussed this problem in detail, but no further nomenclatural act was made. Consequently, the two genera cannot be distinguished reliably using current methods. On the other hand, Nomura and Idris (2005) also noted the similarities between Pseudophanias and the tribe Hybocephalini, but this relationship needs further investigation.
So far, four tmesiphorine genera have been recognized in Japan: Saltisedes Kubota, 1944, Tmesiphorus LeConte, 1849, Tmesiphoromimus Löbl, 1964, and Raphitreus Sharp, 1883(Shibata et al. 2013Yin et al. 2013b;Inoue et al. 2020), while none have been found from Taiwan. In this paper, we formally place Chandleriella, syn. n., as a junior synonym of Pseudophanias and describe three new species from Japan and Taiwan.

Material and methods
The type specimens of Pseudophanias species in the Raffray's Collection were examined by the second author at the Muséum National d'Histoire Naturelle, Paris, France (MNHN). The holotypes of Pseudophanias yaimensis, P. nakanoi and P. excavatus are deposited in the National Museum of Nature and Science, Tokyo, Japan (NSMT) and paratypes are deposited in NSMT, and the Kyushu University Museum, Fukuoka, Japan (KUM). Two paratypes of Pseudophanias yaimensis are temporally deposited in the Insect Collection of Shanghai Normal University, Shanghai, China (SNUC), and will be eventually housed in the National Museum of Natural Science, Taichung, Taiwan (NMNS).
The label data of the holotypes are quoted verbatim. A slash (/) was used to separate lines on the same label, and a double slash (//) was used to separate different labels on the same pin.
The specimens were soaked in distilled water overnight, and male genitalia were obtained by removing tergites and sternites VIII-IX. The male genitalia were soaked in cold 10% KOH for about 6 hours, and afterward they were washed in distilled water for 10 minutes. They were subsequently transferred to 50% ethanol for 2-5 minutes and then to 80% ethanol for 2 minutes. Finally, the male genitalia were soaked in 99% ethanol for 10 minutes and were then mounted in Euparal on a 5 × 10 mm micro-cover glass. The micro-cover glass with male genitalia was glued onto a paper card (5 × 7 mm) and pinned under the specimen (Maruyama 2004). The specimens were also examined using a scanning electron microscope (SEM; Hitachi SU-3500) at the Center for Advanced Instrumental and Educational Supports, Faculty of Agriculture, Kyushu University. For the SEM observations, all examined materials were coated with gold by Ion Sputter (Ion sputter; Hitachi MC1000) and examined at 5.00 kV and 30 Pa.
The following abbreviations were applied: AL length of the dorsally visible part of the abdomen along the midline; AW maximum width of the abdomen; EL length of the elytra along the suture; EW maximum width of the elytra; HL length of the head from the anterior clypeal margin to the occipital constriction; HW width of the head across the eyes; PL length of the pronotum along the midline; PW maximum width of the pronotum.
Length of the body (BL) was a combination of HL + PL + EL + AL. All measurements are recorded in millimeters (mm).  Raffray, 1904, but can be distinguished by the distinctly smaller body size (3.00-3.20 mm in P. robustus), the angulate antennomere 11 at anterolateral margins in the male, the finely punctate head and pronotum, and the shorter discal carinae on tergite IV.
Etymology. Ishigaki Island, where the type locality of this species was discovered, is a part of the Yaeyama Islands. This specific epithet refers to Yaima which is a local dialect of the Yaeyama Islands.
Remarks. This species is distributed in Yaeyama-shotô Islands, Japan and Taiwan, China. The two localities are close to each other and shared the same fauna for some insect groups. The two populations show slight difference in the morphology of the aedeagus. The lateral projections of the median lobe of the population of Taiwan are relatively longer and narrower than those of the population of Yaeyama. But the general appearance and especially the male sexual characters are otherwise almost identical. Therefore, we treat such a difference as interspecific variation.
The Taiwanese specimens were collected from a nest of the Nasutitermes parvonasutus termite. However, the Japanese specimens were collected from leaf litter samples or by FIT. Some pselaphine species are known to live under the bark and rotten wood with termites. Thus, more information is needed to recognize the possible termitophyly of the new species.

Diagnosis.
Pseudophanias nakanoi is similar to P. clavatus Raffray, 1904, but P. nakanoi can be distinguished from the latter by its modified antennal clubs, which are formed by three apical antennomeres, and the finely punctate head and pronotum.
Etymology. The new species is named after Mr Fumitaka Nakano, the original collector of the holotype.
Distribution. Japan (Ryûkyû: Tokara-rettô Isls.; Kyûshû: Yakushima Is.) Biology. The holotype was collected from a dead tree of the family Fagaceae, and the paratype female was collected from leaf litter.

Diagnosis. This species is readily distinguished from other members of
Pseudophanias by the clasping formed antennae in the male, frontal sulcus indistinct, and the rounded pronotum.
Etymology. The specific epithet refers to the strongly excavated antennae in the male of the new species.
Distribution. China (Taiwan). Biology. This species was collected from leaf litter.
Diagnosis. This species is readily distinguished from other members of Pseudophanias by a combination of the following character states: Body length over 3.50 mm; antennal club formed by antennomere 11 alone; antennomere 11 strongly enlarged and modified to form bowl-like in male; aedeagus symmetric, median lobe tri-lobed at apex; parameres elongate, narrowing from base toward apex, with several long apical setae .

Discussion
Members of the supertribe Pselaphitae usually have two equal tarsal claws and have posterior claws smaller than the anterior ones in some genera. Some tribes, such as Pselaphini, have singular tarsal claws (Chandler 2001). The tribe Tyrini typically has two tarsal claws equal in size, but a few genera have protarsi with posterior tarsal claws that are reduced in size (Hlaváč and Chandler 2005). The tribe Tmesiphorini has the claw morphology similar to Tyrini (Chandler 2001), but clearly differs in the setose sulci that embrace the antennal bases. The morphology of the claws is frequently used for classification in some genera. However, interspecies differences in tarsal claw morphology have been recognized in the genus Tmesiphorus (Inoue et al. 2019). Although the genus Phalepsus Westwood of the tribe Phalepsini is distinguished from the other tribes by its strongly asymmetrical tarsal claws (Raffray 1890a;Jeannel 1949), some species have claws that are nearly subequal in size in Phalepsus (Chandler 2001). Therefore, tarsal claw morphology may vary at the species level within a genus in Pselaphitae. The genus Chandleriella was tentatively separated from the genus Pseudophanias by a number of external characters (see Introduction). In this study, the three new species we placed in Pseudophanias show intermediate and different ratios of posterior and anterior tarsal claws. Additionally, in Pseudophanias, many undescribed species are recognized in Southeast Asia, and their posterior tarsal claws are reduced to various degrees in each species (Nomura pers. obs.). Therefore, the genera Pseudophanias and Chandleriella cannot be separated based on their tarsal claw morphology, and Chandleriella, syn. nov., is here synonymized with Pseudophanias. The two recognized species of Chandleriella are here removed to Pseudophanias, resulting in P. termitophilus (Bryant, 1915) comb. nov., and P. yunnanicus  comb. nov.
List of world species on our previous version of the manuscript. We are very grateful to Dr James Hammond (Natural Resources Canada, Alberta) for his critical reading and invaluable comments on our manuscript. The third author Yin was supported by the National Natural Science Foundation of China (No. 31872965), and the 'Phosphor' Science Foundation, Shanghai Municipal Science and Technology Commission, China (19QA1406600).