Research Article |
Corresponding author: Wen-Xuan Bi ( insectb@163.com ) Corresponding author: Mei-Ying Lin ( linmeiying2021@126.com ) Academic editor: Francesco Vitali
© 2016 Wen-Xuan Bi, Mei-Ying Lin.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Bi W-X, Lin M-Y (2016) A revision of the genus Pseudoechthistatus Pic (Coleoptera, Cerambycidae, Lamiinae, Lamiini). ZooKeys 604: 49-85. https://doi.org/10.3897/zookeys.604.9049
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The genus Pseudoechthistatus Pic, 1917 is redefined and revised. Five species of the genus are described as new, P. sinicus sp. n. and P. chiangshunani sp. n. from central Yunnan, China, P. pufujiae sp. n. from western Yunnan, China, and P. holzschuhi sp. n. and P. glabripennis sp. n. from southern Yunnan and northern Vietnam. Pseudoechthistatus birmanicus Breuning, 1942 is excluded from the fauna of China. Three poorly known species, P. obliquefasciatus Pic, 1917, P. granulatus Breuning, 1942, and P. acutipennis Chiang, 1981 are redescribed, and the type localities of the former two species are discussed. Endophallic structure of seven species in inflated and everted condition are studied and compared with their relatives. Illustrations of habitus and major diagnostic features of all species are provided. Some biological notes are reported. An identification key as well as a distributional map are presented.
Taxonomy, new species, Lamiini , endophallus, China, Vietnam, Myanmar, Oriental region
The little-known genus Pseudoechthistatus Pic, 1917 was established based on a flightless species, P. obliquefasciatus Pic, 1917 from Dali, Yunnan, China. Later,
Specimens of the genus Pseudoechthistatus were so rare that all four species were described from single specimens and only a few additional specimens have been reported since the original publications.
In the course of our studies of material from several major collections and from several expeditions to Yunnan, China, during 2010 to 2015, five new species were discovered (including four winged species). The generic definition of Pseudoechthistatus is broadened to legitimately include all those species. The four known species were determined based on high-quality photographs of their type specimens, three of them were reexamined and redescribed based on fresh material. Pseudoechthistatus birmanicus is excluded from the fauna of China, and the type localities of P. obliquefasciatus and P. granulatus are discussed. Endophallic structure of seven species in inflated and everted condition are described, figured and compared with their relatives from Paraleprodera Breuning, 1935. The basic observing method for endophallic comparison is discussed and proposed to be done in everted and inflated condition at least in Lamiinisensu lato.
Specimens are deposited in the following institutions, museums or private collections; abbreviations as shown in the text:
CBWX Collection of Wen-Xuan Bi, Shanghai, China
CCCC Collection of Chang-Chin Chen, Tianjin, China
CCH Collection of Carolus Holzschuh, Villach, Austria
CGQH Collection of Gui-Qiang Huang, Chongqing, China
CHTL Collection of Tian-Long He, Huainan, Anhui, China
CJM Collection of Ming Jin, Shanghai, China
CLB Collection of Bin Liu, Beijing, China
CLC Collection of Chao Li, Beijing, China
CSXB Collection of Xiao-Bin Song, Shanghai, China
CTT Collection of Tomáš Tichý, Opava, Czech Republic
CZDY Collection of De-Yao Zhou, Shanghai, China
MHBU
SHEM
Shanghai Entomology
SWU
College of Plant Protection, Southwest
Labels of the type specimens are quoted verbatim; double quotation marks (“ ”) are used for a single label, a slash (/) is used to separate lines on the same label, italics indicate handwriting, notes are included in [], Chinese characters are transcribed in the modern system.
Terminology of endophallic structures follows
Pseudoechthistatus Pic, 1917: 6. Type species: Pseudoechthistatus obliquefasciatus Pic, 1917, by monotypy.
Pseudechthistatus
(sic):
Pseudoechthistatus
:
Body elongate, medium sized (ca. 15.0–25.0 mm long). Head subequal to the pronotal width at base. Eyes coarsely faceted, strongly emarginate; lower lobe small, weakly prominent, subequal to or slightly longer than width. Frons wider than long. Antennal tubercles moderately prominent and separated. Antennae long, ca. 1.6–2.0 times (in male) or 1.2–1.4 times (in female) as long as body length; scape moderately long, apical cicatrix completed, the 3rd antennomere longest, ca 1.5–1.8 times as long as scape, 4th antennomere slightly longer than (in male) or subequal to (in female) scape, 4th to 10th successively shortened and narrowed, last antennomere slightly longer than penultimate; basal antennomeres (3–4 in male, 4–7 in female) distinctly fringed beneath. Both maxillary and labial terminal palpomeres fusiform. Pronotum cylindrical, subequal to or slightly longer than width at base; with two indistinct transverse grooves at the anterior and posterior margins; disk with a rugose longitudinal ridge, slightly raised medially, both sides with a developed or reduced, longitudinal pubescent band; with a lateral spine moderate long and acute apically at anterior middle of each side; prosternal intercoxal process narrow, slightly emarginate at apex, lower than coxae; procoxal cavities closed posteriorly; mesosternal intercoxal process without tubercle and obliquely sloped in lateral view; mesocoxal cavities open externally to mesepimera; metasternum short to moderately long, ca 1.0–1.8 times as long as mesosternal length. Scutellum broadly rounded posteriorly. Elytra elongate, ca. 1.8–2.2 times as long as humeral width, widest at the middle or at humeri or subparallel-sided in basal half, gradually to strongly narrowed after the middle, rounded or obliquely truncated to acute apically; disk finely to coarsely punctured, granules absent or moderately to strongly developed, with few erect or suberect setae; each elytron conspicuously with a moderate to large sized, median, moderately to strongly raised, glabrous tubercle subbasally (Figs
Male genitalia. Tergite VIII (Figs
Female genitalia. Setae of sternites VIII sparse and short, apical ones longer (Fig.
This genus is unique with a conspicuous raised subbasal tubercle on each elytron among the oriental genera of Lamiini. It is superficially resembles Mesechthistatus Breuning, 1950, but immediately distinguished by antennal scape with a complete cicatrix, basal antennomeres distinctly fringed beneath, pronotum with a rugose median longitudinal ridge, and elytra lacking lateral carinae. Pseudoechthistatus shares some characters with Paraleprodera Breuning, 1935: antennae normal (without swollen), scape with a complete cicatrix, basal antennomeres distinctly fringed beneath, pronotal lateral spine present, prosternal process normal (not angularly enlarged between coxae), protibia with a subapical tooth beneath (at least in male), and similar to some species of Paraleprodera (e.g. Par. diophthalma, Par. bisignata, Par. bigemmata) by possess the subbasal tubercle (or tubercles) on each elytron, but is distinguished by elytron with single large raised subbasal tubercle, pronotum with a rugose longitudinal ridge medially, endophallus with CT developed, swollen in dorsal and ventral sides and APH without a pair of U-shaped sclerite (the latter with the subbasal tubercle composed of small granules, CT of endophallus simple and APH with a pair of U-shaped sclerite (Figs
The subbasal tubercle on each elytron of this genus is usually single and complete. However, a few individuals (two of nearly one hundred specimens) have the subbasal tubercle separated by several grooves (Fig.
The type species of this genus is flightless, having a shortened metasternum (subequal to mesosternum in length), constricted humeri and reduced hindwings. These three related structures were considered as generic characters by
Pseudoechthistatus was placed originally in the tribe Phrissomini by
Pseudoechthistatus
obliquefasciatus
Pic, 1917: 7. Type locality: Tali, Yunnan, China. Type depository:
Pseudechthistatus
(sic) obliquefasciatus:
Pseudoechthistatus
obliquefasciatus
:
Holotype (Fig.
(6 males, 12 females): 1 female, Yunnan Weixi Pantiange, 2900 m, 1981.VII.21, leg. Shu-Yong Wang (
Male (Fig.
Body elongate, oblong oval. Head (Fig.
Male genitalia (Figs
Female (Fig.
Elytron with predominant brown pubescence, middle band strongly oblique, subbasal tubercle as wide as or slightly narrower than scutellar width; elytral apices rounded; humeri slightly constricted; hindwings strongly reduced. Endophallus with APH constrictive, apical bulb (ab) heavily sclerotized apically in ventral side, obliquely truncated in lateral view.
(Map
Slightly intraspecific variation can be observed between the population from northern area (Weixi County: Najieluo; Deqin County: Yanmen) and southern area (Weixi County: Pantiange; Yulong County: Ludian, Judian), the northern population with elytra relatively long: elytral length / humeral width ca. 2.1 in male and 2.3 in female, while in southern population, elytral length / humeral width ca 2.0 in male and 2.2 in female. And the holotype is identical to the population from southern area. Currently, sympatry has not been confirmed among the flightless species of this genus, with the exception that the holotype from “Tali” seemingly overlaps with the range of another flightless species, P. sinicus. However, the old name “Tali” or “Tali Fu” (Breuning, 1942), which had been abandoned in 1913 covered a vast region including today’s Dali City, Yunlong County, Eryuan County, Binchuan County, Xiangyun County etc. (
Pseudoechthistatus acutipennis Chiang, 1981: 80, 84, pl. 1, fig. 7. Type locality: Mt. Omei, Sichuan, China. Type depository: SWU.
Pseudoechthistatus
acutipennis
:
Pseudechthistatus
(sic) acutipennis:
Holotype (Fig.
1 female, Sichuan, Emeishan, Jiulinggang, 1900 m, 2014.VIII.7, leg. De-Yao Zhou (CZDY).
Female (Fig.
Body elongate, oblong oval. Head (Fig.
Male. Unknown.
Body and elytra brownish black, very finely pubescent (besides the tawny pubescent markings); pronotal longitudinal bands reduced, shorter than one-third of pronotal length; elytral middle bands widely interrupted near lateral margin, preapical stripe absent; elytral apices strongly acute, disk densely and coarsely punctured; hindwings reduced.
(Map
Pseudechthistatus
(sic) birmanicus Breuning, 1942: 133. Type locality: Ruby Mines, Myanmar. Type depository:
Pseudoechthistatus
birmanicus
:
Pseudechthistatus
(sic) birmanicus:
Holotype (Fig.
(based on quality photographs, and modified from the original description). Male. Body length 21.0 mm, body width 7.5 mm. Body dark brown, body covered with tawny and brown pubescence. Head with four short tawny vittae behind upper eye lobes. Antennal scape with sparse light yellowish pubescence, basal half of 3rd antennomere with sparse fine light yellowish pubescence. Pronotum with paired discal longitudinal band rather long, longer than two-thirds of pronotal length. Elytron with pubescence predominantly brick-red; middle pubescent band light yellowish, broad, well defined, nearly transverse, reaching suture; the preapical stripe same color as middle band, well developed, moderately broader at base. Body elongate. Antennae 1.7 times as long as body length, surpassing elytral apex by five antennomeres; 3rd antennomere 1.7 times as long as scape, 1.1 times as long as 4th antennomere; scape moderately punctured, 3rd antennomere sparsely punctured on basal half; scape to 3rd antennomere fringed beneath. Pronotum slightly longer than width at base, lateral spine short, slightly thickened at base with moderate acute apex; metasternum 1.8 times as long as mesosternal length. Elytra 1.6 times as wide as pronotal base at humeri, 1.8 times as long as humeral width; subparallel-sided in basal one-fourth, very weakly widened a little before middle, then moderately convergent toward subacute apices; disk sparsely and finely punctured, sparsely provided with large but flat granules extending to apical one-fourth; subbasal tubercle close to elytral base, moderately developed and raised, ca. 1.3 times as wide as scutellar width. Hindwings developed, distinctly longer than elytral length.
(Map
This species is only known from its type locality, Ruby Mines (= Mogok), Myanmar at present. Based on our examination of photos of the holotype, the distribution of this species in Yunnan, reported by
Pseudechthistatus
(sic) granulatus Breuning, 1942: 133. Type locality: Tatsienlu (?). Type depository:
Pseudoechthistatus
granulatus
:
Pseudechthistatus
(sic) granulatus:
Holotype (Fig.
(22 males, 20 females): 1 male, Yunnan Prov., Gaolinggongshan, Fugong County, Shiyueliangxiang, Shibaliyingdi, 3105 m, 27.18380°N, 98.71021°E, 2004.V.7 night, leg. Hong-Bin Liang (
Male. (Fig.
Body elongate, oblong oval. Head (Fig.
Male genitalia (Figs
Female (Figs
Lower eye lobe rather short, 0.5 times as long as gena (in female); elytron with granules large and raised, sparsely scattered throughout, middle band variable, complete or interrupted to nearly interrupted near lateral margin; humeri slightly constricted; hindwings strongly reduced. Endophallus with APH constrictive, apical bulb (ab) heavily sclerotized apically, obliquely truncated in lateral view.
(Map
Based on the morphological similarities, the population from Gongshan County and Fugong County of Yunnan Province are considered as P. granulatus temporarily. The type locality of P. granulatus, “Tatsienlu”(= Kangding County) perceived to be doubtful for the following reasons. Based on reliable collecting data, the distribution of a flightless species (at least among this genus) does not support such remote distance (more than 400 km away from “Tatsienlu” to the population in Yunnan). Moreover, no individual has thus far been found in the intervening area. The population from Gongshan and Fugong could not be separated from the type specimen from “Tatsienlu” morphologically. The weak differences, such as slightly longer antennae and lighter pubescence color should be treated as intraspecific variation. Furthermore, females from Yunnan share the shorter lower eye lobes with the female type specimen, while other congeners have longer lower eye lobes (except for P. pufujinae sp. n.). The poor and handwritten label of the holotype (Fig.
Holotype: male, “Yunnan, Dayao County, Santaixiang / Xiaobaicaoling / 2980 m 2013.V.29–30 / leg. Wen-Xuan Bi” (
Male (Fig.
Habitus of Pseudoechthistatus spp. 21–28 head in frontal view 29 subbasal tubercle of elytron showing an abnormal form 30–37 femora in ventral view showing pubescence patterns 21, 29, 30 P. obliquefasciatus Pic, 1917 22, 31 P. granulatus Breuning, 1942 23, 33 P. glabripennis sp. n. 24, 34 P. sinicus sp. n. 25, 35 P. chiangshunani sp. n. 26, 36 P. holzschuhi sp. n. 27, 37 P. pufujiae sp. n. 28, 32 P. acutipennis Chiang, 1981. a male b female.
Body elongate, oblong oval. Head (Fig.
Habitus of Pseudoechthistatus spp (a male b female). 38–40 basal elytra in dorsal-lateral view showing shape and size of subbasal tubercles 41–48 hindwings of Pseudoechthistatus spp., scale = corresponding elytral length. 38, 41 P. obliquefasciatus Pic, 1917 39, 42 P. granulatus Breuning, 1942 43 P. sinicus sp. n. 44 P. acutipennis Chiang, 1981 45 P. pufujiae sp. n. 46 P. chiangshunani sp. n. 47 P. holzschuhi sp. n. 40, 48 P. glabripennis sp. n.
Male genitalia (Figs
Terminalia of Pseudoechthistatus spp. 49, 56 P. obliquefasciatus Pic, 1917 50 P. granulatus Breuning, 1942 51 P. glabripennis sp. n. 52 P. sinicus sp. n. 53 P. chiangshunani sp. n. 54 P. holzschuhi sp. n. 55 P. pufujiae sp. n. 49–55 male. a tergite VIII with sternites VIII & IX b tegmen in ventral view c ditto in lateral view d median lobe in ventral view e ditto in lateral view 56 female. f sternite VIII g ovipositor in dorsal view h ditto in ventral view. Scale 1 mm.
Terminalia of Pseudoechthistatus and Paraleprodera species. 57 spermatheca 58–67 endophallus in inflated and everted condition, lateral view 57–58 Pseudoechthistatus obliquefasciatus Pic 59 P. sinicus sp. n. 60 P. granulatus Breuning, 1942 61 P. pufujiae sp. n. 62 P. glabripennis sp. n. 63 P. holzschuhi sp. n. 64 P. chiangshunani sp. n. 65 Paraleprodera mesophthalma Bi & Lin, 2012 66 Par. carolina (Fairmaire, 1899) 67 Par. triangularis (Thomson, 1865) 68 Par. d. diophthalma (Pascoe, 1857). a enlargement of spermathecal capsule (SP) b endophallus in inflated and non-everted condition, show internal membrane (im) of apical furrow (af) c, dAPH in ventral view eCT in ventral view f, gAPH in dorsal view.
Female (Fig.
This new species can be distinguished from most of the congeners by elytra comparatively shorter (only 1.8 times as long as humeral width), humeri distinctly constricted in both sexes; hindwings reduced; granules moderate in size near humerus and behind basal one-fourth, weakened near apical one-third; APH of endophallus strongly constricted. It is similar to P. obliquefasciatus by color pattern but can be easily distinguished by elytra relatively shorter in length, with bigger subbasal tubercles, median pubescent band not so oblique and APH of endophallus more strongly sclerited. It is similar to P. granulatus by size of elytral subbasal tubercles but can be separated by shorter elytra, weaker granules, and denser ventral tawny pubescence.
The new species is named after the country of its type locality.
(Map
Holotype: male,“Yunnan Prov., Zhengyuanxian / Jiujiazhen, Qianjiazhai / 2012.V.6 2600 m”, “N24°17.143’ / E101°15.060’ / leg. Ling-Zeng Meng ”(
Male. (Fig.
Body elongate, oblong oval. Head (Fig.
Male genitalia (Figs
Female (Fig.
This new species is similar to P. sinicus sp. n., but is distinguishable by having the hindwings fully developed, punctures on elytra distinct and deeper, and APH of endophallus swollen. It resembles P. holzschuhi but differs in having antennae (at least 3rd to 4th antennomeres) without yellowish annulations, 3rd antennomere fringed beneath (fringed only at basal half in P. holzschuhi), punctures and granules on elytra more developed, elytra wider (compare with its length), humeri constricted in female. It can be distinguished from P. birmanicus by the longer 3rd antennomere, deeper elytral punctures, smaller elytral granules and narrower median band on elytra, and the subbasal tubercle of elytron not so close to elytral base.
The new species is dedicated to the late Shu-Nan Chiang (1914–2013), an entomologist specialized in the taxonomy of Chinese Cerambycidae.
(Map
Holotype: male, “CHINA. Yunnan, Jinping / Fenshuiling / 2311 m 2010.IX.18 / leg. Xiao-Dong Yang” (
Male (Fig.
Body elongate, oblong oval. Head (Fig.
Male genitalia (Figs
Female (Fig.
This new species is most similar to P. birmanicus and P. chiangshunani sp. n. by the general habitus but can be distinguished from the former by the elytral granules being rather weakly developed and limited within basal one-third; elytral punctures finer and sparser; middle band of elytron interrupted or nearly interrupted and more developed pronotal lateral spines. It can also be distinguished from the latter by the antenna being shorter than body length, at least 3rd to 4th antennomeres with light yellowish pubescent annulations at base; elytra relatively smooth, granulate only at basal one-third, elytral punctures finer and sparser; female humeri similar to male, not constricted.
The new species is named after Carolus Holzschuh, a specialist in Cerambycidae, who kindly provided his collection for this study.
(Map
Pseudoechthistatus
birmanicus
:
Holotype: male, “CHINA. Yunnan, Lushui / Yaojiaping 2450 m / 2015.V.4 em. VI.9 / leg. Wen-Xuan Bi”, “IOZ(E)1905345” (
Male (Fig.
Body elongate, oblong oval. Head (Fig.
Male genitalia (Figs
Female (Fig.
This new species can be distinguished from most of the congeners (except P. acutipennis) by elytral disk deeply and coarsely punctured throughout and limited granulated near base. It can be easily distinguished from P. acutipennis by pronotal bands and elytral preapical stripe developed, elytral apices subacute, hindwings developed (in P. acutipennis, pronotal bands reduced, preapical stripe of elytron absent, elytral apices strongly acute and hindwings reduced).
The new species is dedicated to the late Fu-Ji Pu (1932–2002), a specialist in Chinese Cerambycidae.
(Map
Holotype: male, “CHINA. Yunnan / Menglun, 55 km / 650 m 2012.IV.25 / leg. Chao Wu” (
Male. (Fig.
Body elongate, fusiform. Head (Fig.
Male genitalia (Figs
Female (Fig.
This new species is easily distinguishable from congeners by combination of the following characters: elytral disk smooth, very finely punctured and pubescent, with dark purple sheen; middle band of elytron moderately oblique, complete; elytra distinctly widest across humeri (at least in males). Endophallus with the overall shape unique, especially by basal swelling (bs) of CT distinctly tuberculate laterally; APH strongly constrictive and subcylindrical in lateral view.
The new species is named from a combination of the Latin stem, ‘glabri’and ‘pennis’referring to the smooth surface of elytra.
(Map
No biological information has been so far available for Pseudoechthistatus. This overview is based on notes from several collectors and the observation of the first author as well as the label data from the specimens. Most species appear to occur in broadleaf deciduous or mixed coniferous and broadleaf forests (Fig.
Adults were mostly observed on dead leaves and branches: P. granulatus were feeding on dead leaves or bark of Pterocarya sp. (Juglandaceae) and Acer spp. (Aceraceae); the population of P. chiangshunani from Manwanzhen, Lincang City were crawling on the trunk of dead Juglans regia (Juglandaceae) or feeding on dead leaves of Alnus cremastogyne (Betulaceae); some specimens of P. sinicus and P. obliquefasciatus were collected by beating dead branches of Cyclobalanopsis spp. and Quercus spp. (Fagaceae), while the population of P. sinicus in Xiaobaicaoling, Santaixiang, Dayao County, were feeding on living leaves of Acer sp. Some adults of P. pufujiae were reared from larvae collected under bark of a fallen tree of Pterocarya (Figs
Most species are nocturnal, and remain hidden in or around their host plants during daytime (Figs
Besides the species with reduced hindwings which are apparently flightless, P. pufujiae (Fig.
Up to now, only P. glabripennis and P. holzschuhi, both with normally developed hindwings, are known to be sympatric in Fenshuiling, southeast Yunnan (Map
Pseudoechthistatus sinicus and hunting spiders (possibly Lycosidae) were active on ground vegetation simultaneously at night (observed in Weibaoshan and Xiaobaicaoling). In consideration of the elytral subbasal tubercles of Pseudoechthistatus that resemble the posterior median eyes of the spiders, we suppose this resemblance may represent a case of Batesian mimicry, but more evidence is required before any conclusion can be reached.
The shortened metasternum (associated with reduced hindwings) was one of the diagnostic characters used to define Dorcadionini, Morimopsini, Parmenini, and Phrissomini of Lamiinae (
In Breuning’s tribal system of the Lamiinae, the occurrence of species with complete and reduced hindwings within the same genus or subgenus is uncommon, e.g. subgenus Pseudale of Pterolophia (Malihara 1988; Yamasako 2016 pers. comm.) and Spalacopsis (
Investigation of the inflated endophallus in Cerambycidae was considered to have been undertaken for the first time recently (
The previous studies on the inflated endophallus can be subdivided into two paths (
The endophallic terminology of Cerambycidae has been proposed and applied for various taxa by several authors (e.g.
Finally, endophallic structures of twelve species or subspecies of Paraleprodera Breuning, 1935 have been investigated for comparison with Pseudoechthistatus Pic, 1917 in this study. As a result, Pseudoechthistatus can be clearly distinguished from Paraleprodera (see generic diagnosis). However, the endophallic diversity of Paraleprodera is considerable and the genus may be subdivided into at least two groups: the triangularis group containing Par. bigemmata, Par. bisignata, Par. diophthalma with subspecies and Par. triangularis, which are characterized by CT less developed (without a distinct swelling) and APH with a pair of U-shaped sclerites (Figs
1 | Elytral disk smooth, very finely punctured; elytra distinctly widest at humeri (at least in males) | P. glabripennis sp. n. |
– | Elytral disk distinctly punctured and/or granulated; elytra widest near middle or subparallel in basal half | 2 |
2 | Pronotal longitudinal pubescent bands reduced, shorter than one-third of pronotal length; elytron strongly pointed apically, preapical stripe absent | P. acutipennis |
– | Pronotal longitudinal pubescent bands well developed, longer than half of pronotal length; elytron rounded or obliquely truncated to subacute apically, with a more or less distinct preapical stripe | 3 |
3 | Elytral humeri distinctly narrower, elytra widened near middle; hindwings strongly reduced, distinctly shorter than elytral length | 4 |
– | Elytra subparallel in basal half (at least in males); hindwings developed, distinctly longer than elytral length | 6 |
4 | Subbasal tubercle of elytron moderate in size, subequal to or slightly narrower than scutellar width; elytral middle band strongly oblique, more than 40 degrees to transverse axis | P. obliquefasciatus |
– | Subbasal tubercle of elytron large, at least 1.2 times wider than scutellar width; elytral middle band moderately oblique or nearly transverse, less than 30 degrees to transverse axis | 5 |
5 | Elytra long, 2.0 times (in males) or 2.2 times (in females) as long as humeral width; elytral disk sparsely covered with large and raised granules throughout; preapical stripe of elytron reduced, shorter than one-fifth of elytral length | P. granulatus |
– | Elytra comparatively short, 1.8 times (in males) or 2.0 times (in females) as long as humeral width; elytral disk sparsely covered with moderately sized granules which become indistinct near apical one-third, and absent beyond apical one-fourth; preapical stripe of elytron developed, subequal to one-fourth of elytral length | P. sinicus sp. n. |
6 | Elytron distinctly granulated, granules reaching at least apical one-fourth; punctures sparse and shallow | 7 |
– | Elytron weakly granulated, granules reaching at most basal one-third; punctures distinct and deep, at least reaching middle | 8 |
7 | Elytral granules smaller and dense; middle band narrow, moderately oblique, interrupted or nearly so at middle | P. chiangshunani sp. n. |
– | Elytral granules larger and sparse; middle band broad, nearly transverse, not interrupted or narrowed | P. birmanicus |
8 | Elytral punctures moderate, reaching the middle; at least 3rd to 4th antennomeres annulate with light pubescence at base; elytral middle band interrupted or nearly so at middle | P. holzschuhi sp. n. |
– | Elytral punctures deep and coarse, reaching the apex; antennomeres without distinct annular light pubescence; elytral middle band complete | P. pufujiae sp. n. |
We thank Li Chen (SWU), Nobuo Ohbayashi (Miura, Japan), Junsuke Yamasako (Tokyo, Japan), Olivier Montreuil and Gérard Tavakilian (