Research Article |
Corresponding author: Martin Fikáček ( mfikacek@gmail.com ) Academic editor: James Liebherr
© 2020 Fenglong Jia, Zulong Liang, Martin Fikáček.
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:
Jia F, Liang Z, Fikáček M (2020) A review of Himalcercyon stat. nov., with description of a new species from the Chinese Himalaya and an updated key to Asian genera of Megasternini (Coleoptera, Hydrophilidae). Deutsche Entomologische Zeitschrift 67(1): 35-49. https://doi.org/10.3897/dez.67.50078
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Himalcercyon Hebauer, 2002 stat. nov. is elevated to genus rank based on the unique form of its mesoventral elevation. The genus is reviewed, redescribed, and illustrated in detail. Two species are recognized: Himalcercyon mirus (Hebauer, 2002) comb. nov. from Nepal and H. franzi sp. nov. from Chinese Himalaya (Xizang Autonomous Region). Both species are illustrated and diagnosed. An updated key to the Asian genera of the tribe Megasternini (Coleoptera, Hydrophilidae, Sphaeridiinae) is provided, along with the SEM micrographs of ventral morphology of these genera. New replacement name Oreosternum nom. nov. is proposed for Oreocyon Hebauer, 2002 which is preoccupied by Oreocyon Marsh, 1872 (Mammalia, Oxyenidae) and Oreocyon Krumbiegel, 1949 (Mammalia, Canidae).
Asia, morphology, new replacement name, new species, new status, Oriental Region, Sphaeridiinae, taxonomy, Xizang, China
Megasternini is the largest clade of terrestrial water scavenger beetles, containing approximately 580 described species currently classified in 52 genera (
The mountains on the southern margin of the Qinghai-Xizang (Tibetan) Plateau are known for their highly diverse and endemic faunas (e.g.,
Recently, we received a small sample of terrestrial hydrophilids from Motuo County, Xizang Autonomous Region, China, a region in the Himalayas at the southern margin of the Qinghai-Xizang Plateau. In contrast to the more northern regions of the Xizang Autonomous Region, Motuo County includes middle to low elevations and is affected by monsoon rains; it is, therefore, warmer and more humid than the main plateau areas. The material contained a species of the Megasternini which is unique in the morphology of its mesoventral plate. We originally considered it to be an undescribed genus, but a detailed survey of megasternine taxa described from the Himalaya region revealed that Cercyon mirus Hebauer, 2002 from Nepal, which was assigned to the monotypic subgenus Himalcercyon Hebauer, 2002 in the original description (
We examined the type series of Cercyon mirus and the small series (10 specimens) of the new species from Motuo County. Male genitalia of the holotypes of both species were examined and photographed in the original position (i.e. with the median lobe inserted in the tegmen). Due to the very limited material available, separation of the median lobe is not always easy and sometimes results in partial damage of some parts of the aedeagus. Genitalia were photographed in glycerol. The aedeagus of the holotype of C. mirus was subsequently embedded in a drop of alcohol-soluble Euparal resin on a piece of glass glued to a small piece of cardboard attached below the respective specimen. Habitus photographs were taken using a Canon D-550 digital camera with attached Canon MP-E65mm f/2.8 1–5 macro lens. Genitalia were photographed using a Canon D1100 digital camera attached to an Olympus BX41 compound microscope (C. mirus) or using an Olympus SZX7 stereomicroscope (new species); combined, focus-stacked images were made with Helicon Focus (Helicon Soft Ltd, Ukraine) software. Scanning electron micrographs of C. mirus and of the Asian genera of the Megasternini were taken using a Hitachi S-3700N environmental electron microscope at the Department of Paleontology, National Museum in Prague; SEMs of the new species were taken using a Phenom Prox scanning electron microscope in the Biological Museum of the Sun Yat-sen University. Images were combined into figures using Adobe Photoshop CS6. All original images, including additional views not presented in this paper, are included in the dataset submitted to the Zenodo archive (https://zenodo.org/ under https://doi.org/10.5281/zenodo.3693743. SEMs of the megasternine genera for the identification key are mostly based on specimens deposited in
Examined specimens are deposited in the following collections:
Cercyon (Himalcercyon) Hebauer, 2002: 39.
Cercyon (Himalcercyon) mirus Hebauer, 2002.
Dorsal surface pubescent; anterior margin of clypeus rounded; frontoclypeal suture not forming transverse ridge between eyes; eyes small, separated 5–6× the width of one eye; prosternum strongly carinate medially, without ridge demarcating median portion from lateral portions (Figs
Body broadly oval and moderately convex; body outline not interrupted between pronotum and elytra.
Head. Excised in front of eyes laterally, antennal base exposed. Labrum concealed under clypeus, not exposed dorsally. Clypeus not deflexed, truncate anteriorly, without anterolateral extensions; anterior margin narrowly beaded. Frontoclypeal suture obsolete, only visible as impunctate bar. Frons with even surface. Eyes rather small, rounded, projected laterally; interocular distance ca 5–6× the width of one eye in dorsal view. Dorsal punctation of head consisting of punctures each bearing a long seta. Maxillary palpus slightly longer than half of width of head, with ventral sucking disc in male; palpomere 2 strongly swollen, longer than palpomere 3; palpomere 4 symmetrical, slightly shorter than palpomere 2, but longer than palpomere 3. Mentum ca 2.1–2.4× as wide as long, trapezoidal, anterior margin not emarginate medially (Figs
Prothorax.
Pronotum relatively short and transverse, widest at base; surface smooth, punctation consisting of setiferous punctures, all punctures of the same size and shape; transverse series of punctures along posterior margin absent. Prosternum well developed, slightly tectiform, strongly carinate medially, without elevated median portion or ridge demarcating median portion from lateral parts (Figs
Mesothorax.
Mesoventrite fused to mesepisterna, bearing hydrofuge pubescence; median portion abruptly raised in posterior half to form arrowhead-shaped elevation (Figs
Metathorax. Metaventrite moderately raised medially, forming a bare pentagonal area weakly projected anteriorly between mesocoxae (Figs
Legs. Coxae partly with hydrofuge pubescence, mesocoxae moderately separated (Fig.
Abdomen
with five ventrites covered by fine hydrofuge pubescence; ventrite 1 2× as long as ventrite 2, strongly carinate throughout (Fig.
1 | Body broadly oval, elytra combined 1.1× longer than wide (Fig. |
H. mirus (Hebauer, 2002) |
– | Body moderately oval, elytra 1.3× longer than wide (Fig. |
H. franzi sp. nov. |
China, Xizang Autonomous Region, Motuo County, track from Dayandong to Hanmi, 2200–2400 m a.s.l. [GPS ca 29.4283N, 95.0498E].
Holotype: CHINA ● 1 ♂; Xizang, Motuo County, Dayandong-Hanmi; 2200–2400 m a.s.l.; 13 Aug 2005, Tang Liang leg.;
Paratypes: CHINA ● 9; same data as for holotype;
Form and color.
Body size 2.5–2.8 mm (2.6 mm in holotype), body width 1.5–1.7 mm (1.55 mm in holotype), widest at anterior third of elytra, arcuately narrowing posteriad (Fig.
Head.
Clypeus with moderately dense fine setiferous punctures, smooth between punctures. Frons with punctures coarser and somewhat denser than those on clypeus, smooth between punctures. Mentum 2× wider than long, rugose, with dense coarse punctures (Fig.
Thorax.
Pronotum with punctation similar to that on frons, interstices without microsculpture; lateral marginal bead shortly overlapping to anterior margin but not to posterior margin, stopping at posterior angle. Scutellar shield smooth, with three to five punctures. Elytral striae sharply impressed (Figs
Male genitalia.
Middle lobe of abdominal sternite IX wide, shorter than lateral struts (Fig.
The species is named after Dr Franz Hebauer, a German taxonomist of the Hydrophiloidea who recognized and described Himalcercyon as a subgenus of Cercyon.
Only known from the type locality in the eastern Himalaya (Motuo county, Xizang Autonomous Region, China) (Fig.
Cercyon (Himalcercyon) mirus Hebauer 2002: 39.
Nepal, Kathmandu district, Sheopuri Mt., 2100–2300 m a.s.l. [GPS ca 27.816672N, 85.400000E].
Holotype: NEPAL ● 1 ♂; Kathmandu Distr. Sheopuri Mt.; 2100–2300 m a.s.l.; 25 Jun 1988; W. Schawaller leg.;
Paratypes: NEPAL ● 2 ♀♀; same data as for holotype;
Form and color.
Body size 3.1–3.5 mm (3.4 mm in holotype), body width 2.0–2.1 mm (2.0 mm in holotype), widest at anterior third of elytra, weakly narrowing posteriad (Fig.
Head.
Clypeus with moderately dense fine setiferous semicircular punctures, smooth between punctures. Frons with punctures of the same size and density as those on clypeus, smooth between punctures. Mentum 1.4× wider than long, rugose, with dense punctures (Fig.
Thorax.
Pronotum with punctation similar to that on frons, interstices without microsculpture; lateral marginal bead shortly overlapping to anterior margin but not to posterior margin, stopping at posterior angle. Scutellar shield smooth, with five to seven punctures. Elytral striae sharply impressed (Fig.
Male genitalia.
Middle lobe of abdominal sternite IX narrow, shorter than lateral struts (Fig.
Known from two localities in central Nepal (Fig.
The following key is mainly based in the ventral characters, namely the form of prosternum and meso- and metaventrite, which are illustrated in Figures
1 | Antennal grooves large, reaching to the lateral margin of hypomeron (Fig. |
2 |
– | Antennal grooves absent or small, not reaching to the lateral margin of the hypomeron (Figs |
5 |
2 | Metaventrite with complete femoral lines reaching from posteriomesal portion to anterolateral corner (Fig. |
3 |
– | Metaventrite without complete femoral lines, at most with short vestiges anterolaterally. (Fig. |
4 |
3 | Mesoventral plate wider than long. Prosternum with wide plate without median carina (Fig. |
Cryptopleurum Mulsant |
– | Mesoventral plate approximatley as long as wide. Prosternal plate approximately as long as wide, with more or less distinct median carina (Fig. |
Pachysternum Motschulsky |
4 | Median portion of prosternum roof-like, high (Fig. |
Pacrillum d'Orchymont |
– | Median portion of prosternum with flat hexagonal plate, not carinate medially (Fig. |
Megasternum Mulsant |
5 | Metaventrite with postcoxal ridge widely diverging from posterior margin of coxal cavity and forming an arcuate ridge reaching lateral margin of metaventrite (Figs |
6 |
– | Metaventrite with postcoxal ridge parallel to posterior margin of coxal cavity or nearly so, reaching anterolateral corner of metaventrite and not forming any arcuate ridge (Figs |
9 |
6 | Metaventrite with complete femoral lines crossing the arcuate postcoxal ridge and X-shape in form (Fig. |
Peltocercyon d'Orchymont |
– | Metaventrite without X-shaped structure, femoral lines absent or short, not crossing with arcuate postcoxal ridge (Figs |
7 |
7 | Mesoventral plate widely contacting metaventrite (Fig. |
8 |
– | Mesoventral plate separated from metaventrite by a wide deep gap (Fig. |
Armostus Sharp |
8 | Metaventrite with deep triangular impression along its lateral margin (Fig. |
Morastus d'Orchymont |
– | Metaventrite without such impression (Fig. |
Oosternum Sharp |
9 | Median portion of prosternum highly elevated and/or delimited from lateral portions by sharp ridges (Figs |
10 |
– | Median portion of prosternum finely carinate, not delimited from lateral portions (Figs |
18 |
10 | Pronotum with deep longitudinal grooves (Fig. |
Emmidolium d'Orchymont |
– | Surface of pronotum without distinct longitudinal depressions. Bare portion of metaventrite confined to medial part only. Tiny to moderately large beetles | 11 |
11 | Median portion of prosternum in form of very small triangular, very highly elevated projection. Antennal grooves absent (Fig. |
Chimaerocyon Fikáček, Maruyama, Vondráček & Short |
– | Median portion of prosternum never so tiny and not so highly elevated. Antennal grooves present, even though sometimes rather small. Abdomen never with apical emargination | 12 |
12 | Prosternal elevation with lateral margins deeply excised (Fig. |
13 |
– | Prosternal elevation with lateral margins or ridges straight (Fig. |
14 |
13 | Tiny species, 1.2–1.6 mm. Metaventrite with complete femoral lines (Fig. |
Paroosternum Scott |
– | Large species, ca 3.0 mm. Metaventrite without femoral lines (Fig. |
Oreosternum nom. nov. |
14 | Elytral series deeply impressed with the impressions contiguous to anterior margin of each elytron (Fig. |
15 |
– | Elytral series not impressed or impressions of elytral striae series not reaching anterior margin of each elytron. Mesoventral elevation elongate or as long as wide | 16 |
15 | Pronotum highly bulged in lateral view, not forming a continuous curve with elytra. Anterior margin of prosternal elevation strongly projecting anteriad (Fig. |
Bolbonotum Hansen |
– | Pronotum not highly bulged in lateral view, forming a continuous curve with elytra. Anterior margin of prosternal elevation straight (Fig. |
Kahanga Hansen |
16 | Grooves for reception of procoxae ending far before the anterior margin of mesocoxal cavities (Fig. |
Gillisius d'Orchymont (part)* |
– | Grooves for reception of proxocae reaching nearly the mesocoxal cavities (Fig. |
17 |
17 | Mesoventral elevation nearly semi-elliptical (Fig. |
Australocyon Hansen |
– | Mesoventral elevation more less pentagonal, without any marginal rim (Fig. |
Nipponocercyon Satô |
18 | Abdominal ventrite 1 without median carina. Mesoventral elevation narrowly laminar (Fig. |
Cycreon d'Orchymont |
– | Abdominal ventrite 1 carinate medially. Mesoventral elevation in form of a lamina or an elongate plate | 19 |
19 | Ventral face of meso- and metatibiae with dense, long pubescence. Ventral morphology similar to Figure |
Pilocnema Hansen |
– | Ventral face of meso- and metatibiae never densely pubescent, at most with sparse short setae. Ventral morphology similar to Figures |
20 |
20 | Mesoventral elevation laminar or forming an oval elongate plate; posterior part of the plate rounded or acute (as in Fig. |
21 |
– | Mesoventral elevation elongate, but sharply cut off posteriorly, contacting metaventrite more or less in a straight line (as in Figs |
22 |
21 | Median portion of prosternum with a pair of transverse ridges partly delimiting prosternal process (Fig. |
Pseudocercyon d'Orchymont |
– | Median portion of prosternum without such ridges, only simply carinate (Fig. |
Cercyon Leach |
22 | Mesoventral elevation arrowhead-shaped, with lateral angulate lobes (Figs |
Himalcercyon Hebauer |
– | Mesoventral elevation elongate oval (as in Fig. |
23 |
23 | India, continental Southeast Asia and China. | Gillisius d'Orchymont (part) |
– | Islands of the Malay Archipelago. | Pelosoma Mulsant* |
List of Eastern Palaearctic and Oriental general of the Megasternini, with number of described species and references to the most important keys or taxonomic treatments.
Genus | Described species | Keys or original descriptions |
---|---|---|
Armostus | 11 |
|
Australocyon | 7 |
|
Bolbonotum | 3 |
|
Cercyon | 148 |
|
Chimaerocyon | 2 |
|
Cryptopleurum | 7 |
|
Cycreon | 4 + 1 ssp. |
|
Emmidolium | 1 |
|
Gillisius | 2 |
|
Himalcercyon | 2 | this paper |
Kahanga | 1 |
|
Megasternum | 4 |
|
Morastus | 1 |
|
Nipponocercyon | 3 |
|
Oosternum | 9 |
|
Oreosternum | 1 | Hebauer 2002 |
Pachysternum | 11 |
|
Pacrillum | 5 |
|
Paroosternum | 5 |
|
Pelosoma | 2 |
|
Peltocercyon | 4 |
|
Pilocnema | 1 |
|
Pseudocercyon | 1 |
|
= Oreocyon
Oreocyon frigidus Hebauer, 2002 (= Oreosternum frigidum comb. nov.)
While preparing the key, we noticed that the genus name Oreocyon is preoccupied by two older names: Oreocyon Marsh, 1872 (a fossil oxyaenid mammal, today a synonym of Patriofelis Leidy, 1872) and Oreocyon Krumbiegel, 1949 (a genus of Canidae described based on fur remains, later renamed to Dasycyon Krumbiegel, 1953 due to homonymy and today considered as a synonym of Canis Linnaeus, 1758). To avoid the homonymy, we are here proposing a new replacement name Oreosternum nom. nov. for Oreocyon Hebauer, 2002. The new name combines the prefix oreo- referring to mountains as used in the original name, and the core sternum, referring to the expected close relationship of this genus to Paroosternum Scott, 1913 exhibited by the prosternal morphology (see the key above). The new name is gender neutral.
Ventral view of thorax and additional diagnostic characters of eastern Palaearctic and Oriental genera of the Megasternini. A–C. Ventral view: A. Pseudocercyon andrewesi. B. Cercyon sp. C. Gillisius madurensis, holotype. D–H. Other characters: D. Cryptopleurum coomani, mentum. E. Emmidolium excavatum, pronotum. F. Bolbonotum sp., base of elytra. G. Kahanga inconspicua, holotype, base of elytra. H. Australocyon sp. (A. pilocnemoides group), hind femur.
The genus-level systematics of the tribe Megasternini are currently based on the traditionally understood genera, defined by characters of the prosternum and meso- and metaventrite, i.e. structures which are morphologically very diverse within the clade. Following this approach, it is possible to define small and morphologically rather uniform genera for roughly half of the known species. On the other hand, the remaining half of megasternine species (i.e. ca 270 species) is assigned to the genus Cercyon Leach, 1817 as they are rather uniform in ventral characters. Eleven subgenera are defined inside of Cercyon to facilitate the identification of species, some of which seem to truly group related species (e.g., Arcocercyon Hebauer, 2003, Paracycreon d’Orchymont, 1942), but others very likely grouping unrelated species sharing a single derived character (e.g., Acycreon d’Orchymont, 1942; see
We thank Dr Li-zhen Li, Dr Liang Tang, Mr Jia-yao Hu, Mr Xiao-bin Song (all Shanghai Normal University, China) for their kind help during the visit of the senior author and for the donation of type specimens to us. We are obliged to Max Barclay Christine Taylor (Natural History Museum, London, UK) and Pol Limbourg (Institute Royal des Sciences Naturelles, Brussels, Belgium) for allowing us to examine holotypes of Kahanga inconspicua and Gillisius madurensis whose SEMs we used in the key. We are indebted to B. Clarkson (Instituto Oswaldo Cruz – Fiocruz, Rio de Janeiro, Brazil), S.K. Ryndevich (Baranovichy State College, Baranovichy), A.E.Z. Short (University of Kansas, Lawrence, USA) and J.K. Liebherr (Cornell University, Ithaca, USA) for their comments and corrections of the early draft of this paper. This study was supported by National Natural Science Foundation of China (31772494) to F.-L. Jia and by the Ministry of Culture of the Czech Republic (DKRVO 2019-2023/5.I.b, National Museum, 00023272) to M. Fikáček.