To be or not to be a synonym – revision of the Donaciaclavareaui-fukiensis complex (Coleoptera, Chrysomelidae, Donaciinae)

Abstract The East Palaearctic species Donaciaclavareaui Jacobson, 1906 and Donaciafukiensis Goecke, 1944 have been confused for decades. Finally, D.fukiensis was synonymized with D.clavareaui by Askevold (1990) but he could not examine the type series of D.fukiensis because it was stored in an inaccessible collection. Cong and Yu (1997) re-established D.fukiensis as a distinct species, also without direct access to the type series. The synonymization by Askevold (1990) was applied in the identification key of Palaearctic Chrysomelidae (Warchalowski 2010) and the Catalogue of Palaearctic Chrysomelidae (Silfverberg 2010). Because the type series of D.fukiensis is now accessible, it has been possible to proof that D.fukiensis is a distinct species, and a lectotype has been established from the series of seven syntypes. Donaciakweilina Chen, 1966 and D.mediohirsuta Chen, 1966, which were split from the mixture of D.clavareaui and D.fukiensis, are now also synonymized with D.clavareaui, because their characters are the same or within the variation range of the characters of D.clavareaui. Furthermore, a distribution map is provided with the reliable records known to date.


Donacia kweilina
The type specimens are kept in ASIZ except for two paratypes in ISAC. Chen, 1966 Type locality. China: Yunnan, Shishong-Baana (Xishuangbanna), 22°1'N, 100°48'E, 1200 m a.s.l. Holotype. ♀ "Yunnan: Shishong-Baana, 15.5.1958" The type specimen is retained in ASIZ. The characters of the type specimens of D. kweilina and D. mediohirsuta are analysed by the detailed first description of Chen (1966) and by further character descriptions mentioned in Cong and Yu (1997), who had examined these type specimens.

Species record list
In Table 1 all records of these four Donacia species known to date are listed. The specimens indicated with "det. E. Geiser" or "vid. E. Geiser" were examined. Jacobson (1906) described the species D. clavareaui from Kjachta (Russia) in southeast Siberia. It could be easily distinguished from all other Donacia species known by its pubescent pronotum combined with glabrous elytra. In the subsequent decades several Donacia specimens from East Asia where identified as Donacia clavareaui.

Taxonomic history
In the 1940s Goecke, a world-renowned Donaciinae specialist, examined specimens of D. clavareaui in the collection of the Museum Alexander Koenig in Bonn (Germany). He recognized that the specimens from Fujian (south-east China) were different in some characters which are typical for species limitation in Donacia. In 1944 Goecke published the description of the new species D. fukiensis which he split from D. clavareaui.
The description of Jacobson (1906) as well as the description of Goecke (1944) are both very detailed. However, Goecke did not describe which were the critical different characters for the distinction of D. fukiensis from D. clavareaui. He also published no identification key. Both descriptions match with both species (see Appendix 1, 2). This resulted in many misidentifications of East Asian specimens.
In 1961 Gressitt and Kimoto published their comprehensive volume "The Chrysomelidae of China and Korea". Because there were so many Chinese specimens misidentified as D. fukiensis they assumed that D. clavareaui was restricted to Siberia. Therefore their identification key contains only D. fukiensis. The characters they mention in their key are applicable to both species. Their key became famous and widespread. Subsequently almost all specimens of D. clavareaui outside Siberia were identified as D. fukiensis from then on. Chen (1966) recognized that within D. fukiensis, some specimens have different characters. He split two new species, D. kweilina and D. mediohirsuta, off from what was actually still a mixture of the two species D. clavareaui and D. fukiensis.
In the 1980s Askevold worked on his comprehensive revision of the genus Donacia. He investigated the type specimen of D. clavareaui which has been stored in the collection of the MNHN Paris. He also intended to investigate the type specimen of D. fukiensis stored in the collection Goecke which was then part of the private Coleopterea Museum Frey in Tutzing, Bavaria. Due to the special situation of the Museum Frey (see next chapter) no research on type or other specimens was possible at that time. Therefore Askevold studied series of D. fukiensis from Japan and China, which in fact were D. clavareaui. He concluded that there are no differences to the type specimen of D. clavareaui (he was right!) and therefore erroneously synonymized D. fukiensis with D. clavareaui. In 1990 Askevold published his comprehensive revision of the genus Donacia which has been widely used as a reference since.
In the 1990s Cong and Yu worked on a list of the Donaciinae of China. They recognized some differences in the specimens labelled D. clavareaui from Fujian as compared with specimens from other parts of China (as Goecke did more than 50 years before). Therefore they intended to study the type specimens of D. fukiensis from Goecke in Museum Frey. At that time, once again no loan of specimens was possible, but for a short period during the quarrels about the Frey collection it was stored at the ZSMC (see next chapter). Martin Baehr, the curator of Coleoptera section in Munich was in charge; Cong and Yu wrote to Baehr and asked him to check some critical characters at the syntype specimens of D. fukiensis, and Baehr confirmed these characters. Cong and Yu (1997) therefore removed D. fukiensis from synonymy and published the first identification key to distinguish D. clavareaui and D. fukiensis; they also included D. kweilina Chen, 1966 andD. mediohirsuta Chen, 1966. They also published accurate distribution data of these four species as far as they were substantiated.
The third volume of Water Beetles of China was published by Jäch and Ji in 2003 with Konstantinov as the author of the chapter about aquatic Chrysomelidae (Konstantinov 2003). He refers to all four species mentioned above, but he compiled their distribution data from sources where D. clavareaui and D. fukiensis were confused, and so they are not reliable.
In 2010 two very important comprehensive studies on Chrysomelidae were published: the Identification Key of Palaearctic Chrysomelidae (Warchalowski 2010) and the sixth volume of the Catalogue of Palaearctic Coleoptera which contained the Chrysomelidae in which Silfverberg was the author of the chapter on the Donaciinae (Silfverberg 2010). Both books are very useful and are the results of enormous workloads of the authors. Warchalowski is a specialist for Alticini (Galerucinae, Chrysomelidae) and Silfverberg is a specialist for Criocerinae und Galerucinae. Both wrote the Donaciinae chapter as no Donaciinae specialist was available and they both referred to the last comprehensive work on Donaciinae (Askevold 1990); therefore D. fukiensis is treated as a synonym to D. clavareaui in both volumes.
In 2015 a global checklist on Donaciinae was published (Geiser 2015), based on Silfverberg (2010) for the Palaearctic species and D. fukiensis is treated as a synonym to D. clavareaui there, also.
In 2017 I visited the collection of the SDEI in Müncheberg, Germany, which contains specimens of D. clavareaui and D. fukiensis, both identified by Goecke in 1952. I saw immediately what Goecke and Cong and Yu had seen before: that these two specimens differ in characters which are typical for separate species of Donaciinae. Fortunately the type specimens are accessible now in the NHMB and it was possible to check the characters of the seven syntypes and to finally designate a lectotype.

The Museum Georg Frey and its unusual situation from 1976 to 1997
Georg Frey  was the owner of a clothes-producing company ("Lodenfrey"). He had an ardent interest in beetles, and attended and paid for field trips worldwide to collect beetles; he also bought collections from specialists. Near his house in Tutzing (south of Munich, Bavaria, Germany) he established a private museum and employed up to five scientists and assistants. When the Donaciinae specialist Hans Goecke died in 1963 Georg Frey bought his famous collection containing many type specimens (Anonymous 1963, Evers 1963. In the decades after the WWII scientific institutions like natural history museums had insufficient and often only provisional storage facilities. At the Museum Frey the Goecke collection was well maintained as Frey employed the then-Chrysomelidae specialists, Jan Bechyne and Gerhard Scherer. When Georg Frey died in 1976, a quarrel began in the Frey family. The sons of Georg Frey intended to donate the whole collection to the ZSMC, because that had been the will of their father they argued; but the widow of Georg Frey began negotiations and finally sold the whole collection to the Natural History Museum of Basel, Switzerland. This started a conflict which involved the Frey family, the Munich State collection, several Switzerland institutions, and German Government institutions. The latter declared this beetle collection a national treasure which must not be transferred outside the borders of Germany. In 1992 the widow died and the collection was clandestinely transferred to the ZSMC before the Basel Museum received information on her death. The legal dispute continued and from 1995 onwards the collection was stored in boxes in Weil am Rhein, Germany, a city near Basel at the Swiss border (Furth 1996). In 1997 it was confirmed that the Museum Basel was the legitimate owner of this beetle collection and it was then transferred there (see further details from the Basel perspective in "Käfer für Basel" [https://kaeferfuerbasel.ch/die-sammlung-georg-frey/]). These incidents were the reason that between 1976 and 1998 it was impossible for long periods to borrow specimens and even to visit the collection to examine it in situ. Jacobson (1906)  The head, antennae, legs, and pronota are very similar, but their elytra are strikingly different. The main character differences are -Shape of the contour of the elytra -Punctures of the elytra -Elytral epipleura -Elytral apex -Female: last sternite -Male: aedeagus Table 2. Common and different characters of Donacia clavareaui and Donacia fukiensis. Each character was based on specimens indicated in Table 1.

D. clavareaui D. fukiensis General
Medium sized, pitchy brown, dark bronze, shiny, antennae and legs partially reddish, hind femora don't reach the apex of the elytra, hind femora claviform with acute tooth, pronotal disc with very fine hairs, elytra glabrous Body Shape Habitus like typical Donacia (Fig. 4) Habitus resembles Plateumaris ( Fig. 1) Sex difference Males in general more slender and shorter than the females Colour Dark metallic-bronze, greenish-bronze, metallic-cupreous

Shiny bronze
Colour of antennae and legs Antennae and legs partially yellow, reddish or brown, the extent of the colour is very variable within specimens Ventral Ventral hairs as usual on Donacia, density variable, the colour of the hairs depends on the lighting Size ♂ 6.5-8.0 mm (avg: 7.5), ♀ 8.0-9.0 mm (avg: 8.5) Head Antennae lenght Filiform, slender, almost half as long as the length of the body, in some male specimens reaching farther than the middle of the elytra Antennomeres A2+A3 ≈ A1 ≈ A4 ≈ A5; A2 < A3 The length relations of the single segments to each other are quite variable. The basal parts of the antennomeres are rufous or yellow, the apical parts are dark and sometimes metallic, the ratio between the two colour parts shows a great variation among the specimens Antennal tubercles The antennal tubercles are flattened, with a narrow groove between them Head disc Head disc straight at front with a deep middle groove Calli Calli indistinct, some specimens without calli Frons and eyes Eyes wide apart, the frons width is four times the measured value of the eye width, with no difference between male and female specimens Pronotum Surface Pronotum pubescent, with very fine hairs, on some specimens very difficult to be seen Surface Pronotum finely and densely punctured (Fig. 5) Irregularly punctured, in between the punctures shiny.
Often the punctures are more dense in the anterior and posterior part than in the middle part. Density of the punctures shows a great variation between individual specimens (Fig. 6, 7) Shape Almost quadratic, in some male specimens slightly longer than wide, in some female specimens wider than long. Anterior margin slightly convex, anterior angles well developed, anterior tubercles rather flat, only slightly protruding Scutellum Scutellum with thin and short hairs Elytra Shape Typically Donacia-shaped Rather Plateumaris-shaped General features Approx. twice as long as wide, in most male specimens slightly longer than double width (ratio 2.1), in most female specimens slightly shorter (ratio 1.9) glabrous and shiny Impressions Slightly visible only on some specimens Punctures and intervals Punctures strong and deep, intervals distincly wrinkled (Fig. 8) Punctures very delicate, not deep, intervals only slightly wrinkled, very smooth ( Fig. 9) interval ≈ 4x -7x puncture diameter Epipleura Elytral epipleura approx. as wide or wider than 10 th interval ( Fig. 10) Epipleuron : Interval = 1 : (>) 1 Elytral epipleura narrower than 10 th interval (Fig. 11) Epipleuron : Interval = 1 : (1.5 -2) Apex Elytral apex truncated, the external angle slightly rounded (Fig. 12) Elytral apex indistinctly truncated, evenly and widely rounded with very smooth outer and inner angles ( Fig. 13) Abdomen Pygidium Distinctly arcuately emarginate Truncated and slightly recessed in the middle Male last sternite Apex rectangularly truncated and triangularly impressed Slightly impressed at the apical ridge Female last sternite Basic contour distinctive triangular (Fig. 14) Basic contour convex without a distinctive peak and broadly rounded (Fig. 15)

D. clavareaui D. fukiensis Legs General
Strong legs, all femora clavate, especially at the ♂, at the ♀ mostly more slender, hind femora short, even at the ♂ they don't reach the apex of the elytra by far. Posterior femora with a prominent tooth, which is often broader at the ♂, at the ♀ more slender and more acute. Legs partly reddish, some specimens with completely red anterior tibia, some specimens with rather dark legs Anterior Tibia Anterior tibia shows a protruding tooth towards outward at the insertion of the tarsomere. D. fukiensis: Fig. 18 D. clavareaui: Fig. 4 and https://science.mnhn.fr/institution/mnhn/collection/ec/item/ ec2130?listIndex=1&listCount=6 [26.11.2018] It is clearly visible on most specimens, but on some indistinctly Tarsomeres The 1 st and 3 rd tarsomere have approx. the same length, the 2 nd one is by a third shorter Aedeagus Shape Aedeagus very straight, outer contours in frontal view rather parallel. Median lobe distinctly protruding: Fig. 19, 20, 21 Aedeagus more curved, thickened, narrowed towards the apex. Median lobe slightly protruding: Fig. 22, 23, 24 All these character differences are typical for species in the genus Donacia. There are some well-established species in Donacia which differ in much more subtle characters. Therefore it was correct that Cong and Yu (1997) re-established D. fukiensis as a valid species. Now that the type series of Goecke is available to scientists, I was able to designate a lectotype from the seven syntypes on which the description of Goecke had been based (Fig. 1). Chen (1966) described D. kweilina and D. mediohirsuta which he separated from the mixture of D. fukiensis and D. clavareaui. The common character of these four taxa is the pubescent pronotum combined with glabrous elytra. The first description is published  in Chinese and in English. For practical considerations only the English text is shown in Appendix 3 (for D. kweilina) and Appendix 4 (for D. mediohirsuta). Donacia kweilina is known only from the type series (Cong and Yu 1997). No further records are known.

Character analysis of Donacia kweilina
In Table 3 the characters of D. kweilina are listed according to the original description by Chen (1966) and provided by Cong and Yu (1997), who examined the type specimens. My comments result from the examination of specimens of D. clavareaui.
The characters which should distinguish D. kweilina from D. clavareaui are either the same or within the variations range of D. clavareaui. Therefore D. kweilina is a synonym of D. clavareaui.

Character analysis of Donacia mediohirsuta
Donacia mediohirsuta is known only by the type specimen, a single female specimen from Yunnan, Shishong-Baana (Cong and Yu 1997). No further records are known. In Table 4 the characters of D. mediohirsuta are listed according to the original description by Chen (1966) and supplemented by Cong and Yu (1997), who have examined the type specimen.
According to Cong and Yu (1997) this specimen resembles D. kweilina with only minor morphological differences. As shown in Table 4 the characters are identical or within the range of D. clavareaui. Therefore D. mediohirsuta is also a synonym of D. clavareaui.

Characters of D. kweilina
Comments Colour aeneo-cupreous (♂, ♀) sometimes sky-blue (♂) D. clavareaui is also aeneo-cupreous, sometimes blue males occur in Donaciinae species Antennae and legs entirely deep coloured, not partly rufous This occurs also in other Donacia species where most of the specimens have partially rufous antennae and legs; colour also very variable in D. clavareaui Antennae: third segment slightly longer than second and distinctly shorter than fourth same proportions of antennomeres in D. clavareaui Head with four weak tubercles, the median longitudinal furrow deep and complete. Pronotum more thickly pubescent, very closely punctured, and covered with silvery hairs, the antero-lateral tubercles distinct, the angles fairly strongly produced. Elytra rather smooth on inner disc, the punctures oblong, the interstices broad, approx. 2-3 times as broad as the cross diameter of the punctures. Apex truncate with the outer angles broadly rounded.
All these characters can be clearly seen at the holotype specimen of D. clavareaui Elytral epipleuron narrow and divided from outermost interval by sharp ridge throughout the entire length of elytra This character is also clearly shown at D. clavareaui (Fig. 10) Last abdominal segment of ♀ much longer and somewhat triangular in shape (Fig. 16) Same typical shape as D. clavareaui (Fig. 14) Hind femora (♂, ♀) broadly toothed beneath, the femora of ♂ not distinctly thicker than those of ♀ Same as D. clavareaui, thickness of hind femora variable Aedeagus: Apex of median lobe cordiform (Cong and Yu 1997) Cong and Yu (1997)   Third antennae segment distinctly longer than the second one, but slightly shorter than the fourth one Same as D. clavareaui

Pronotum more transversal
In D. clavareaui the pronotum is as long as wide or slightly longer than wide; female specimens of Donacia sp. sometimes have a slightly broader pronotum Pronotum finely pubescent only on the median groove Pronotum pubescence varies in D. clavareaui The longitudinal furrow of interocular area much deeper, extending uninterrupted to between the supraantennal tubercles These characters are distinctly visible at the holotype specimen of D. clavareaui Anterior tibiae scarcely produced at apex Variable; the protruded angle of the anterior tibia is mostly distinct, but in some specimens difficult to recognize Hind femora (♀) very weekly toothed beneath Variable in Donacia sp., especially female specimens have weak teeth in comparison with male specimens Last abdominal sternite (♀) more strongly angulate at apex (Fig. 17) Same typical shape as D. clavareaui (Fig. 14 Punctures on elytra rather strong, intervals one to two (sometimes three) times as wide as the diameter of the punctures, elytral epipleuron approx. as wide or wider than 10 th interval, elytral apex truncate (Fig. 12), the angles slightly rounded, female last sternite broadly triangular with posterior margin projected (Fig. 14), aedeagus rather straight and the median lobe cordiform with apex abruptly pointed (Figs 19, 20, 21 Punctures on elytra rather fine, intervals three to seven times as wide as the diameter of the punctures, elytral epipleuron less wide (ca. ½ or ¾ of width) than 10 th interval, elytral apex rounded (Fig. 13), female last sternite broadly rounded (Fig. 15), aedeagus curved and the median lobe with slightly protruding apex (Figs 22, 23, 24  and east of 100° longitude. It is obvious that D. clavareaui must occur in many more locations than those shown in Fig. 25.
Donacia specimens are difficult to collect. The adults can be caught only during a period of a few weeks in late spring and early summer. This period shifts every year due to local weather conditions. Most rare species are found within groups of many specimens of other similar looking, more common Donacia species, and they are therefore often overlooked.

Ecology
All Donaciinae species develop and feed on plants associated with water. As far as the food plants are known, Donacia species are monophagous or oligophagous. Some adults feed on pollen, mostly on Cyperaceae (Kleinschmidt and Kölsch 2011).The larvae live attached to the roots in the sediment. They breathe by piercing the aerenchyme of their food plant with two hollow abdominal stilettos, which are connected to their tracheal system.
The larva of D. clavareaui has been described by Narita (1991Narita ( , 2003. The specimens were collected from roots of the Cyperaceae species Scirpus fluviatilis (Torr.) in Ibariki-ken in Honshu, Japan. According to Bienkowski (2014)

Discussion
If specimens of D. clavareaui and D. fukiensis are compared directly, the differences are striking, especially of the elytra. Although the first descriptions of these species are comprehensive and detailed, they both described both species. Furthermore, it was not possible to create a reliable identification key without correctly identified specimens to hand. This created a vicious circle and caused decades of misidentifications, as well as the splitting of new species from a conglomerate of what was in fact two species. The situation was worsened by the inaccessibility of the type series of D. fukiensis in the Frey collection for a long period.
If specimens are identified incorrectly, all further studies on ecology and distribution are useless. In Figure 25 only reliable data of correctly identified specimens are used. In fact, it shows more the serendipity of the collectors than the reality of the distribution, but this is always the case within rare species. There are certainly more specimens stored in collections throughout the world, but they need to be examined and re-identified in light of the current classification as they may have been mistaken for other Donacia species. Donacia fukiensis may be also hidden within specimens of Plateumaris.
It is also very difficult to infer the distribution of D. clavareaui from its food plant. According to GBIF [https://www.gbif.org/species/2718286; 24.10.2018] Scirpus fluviatilis occurs outside of North America only in Japan and Korea and some spots on the east coast of Australia. The data provided by KewScience [https://wcsp.science.kew. org/namedetail.do?name_id=221898; 24.10.2018] indicate further records from New Zealand, but no records in Asia; GBIF shows only one record of Isolepis fluitans from Ceylon. Scirpus maritimus is widespread, but there is only one record from China and none from Russia. It is very likely that D. clavareaui feeds on Scirpus sp. sensu lato.
Although both species are rare, I hope this paper will trigger some interest to examine the fauna more carefully during field trips in this area. If recent sample sites are known, it would be possible to find the food plant and larvae of D. fukiensis and to analyse the DNA of both species, to include them in the phylogenetic tree published by Kölsch and Pedersen (2008). Because the development of a pubescent upper side occurred several times in the evolution of the genus Donacia it is likely that they are not closely related. some good advice as a reviewer. Martin Baehr (ZSMC) and Michael Theo Schmitt (Ernst-Moritz-Arndt University Greifswald, Germany) provided useful information about the G Frey and J Klapperich collections. Furthermore, I would like to thank Michaela Brojer, Harald Bruckner (both NHMW), and Remigius Geiser jun. (Vienna, Austria) for their support with the photographs and image processing. Remigius Geiser sen. (Salzburg, Austria) translated the Latin text of Jacobson (1906) into German and helped to identify locations in China. I would like to thank him also for correcting the manuscript and helpful discussions. I also benefitted from the helpful advice of the reviewer Horst Kippenberg (Herzogenaurach, Germany).
metallic-cupreous, rufous are the basal parts of all antennae segments ([and] the major part of the apical ones), the whole palpae, the apical part of the mandibles, the apical margin of the labrum, the trochanters, the basal third and the end part of the femora, all tibiae and almost all [on the upper part slightly brownish] tarsomeres. Head Caput oculis sat magnis valdeque prominentibus; The head with quite large and very protruding eyes; temporibus dense scopariis; the tempora with dense, brush-like hairs; canaliculo mediano profundo latoque; the middle groove deep and broad; tuberculis frontalibus indistinctis.
the frontal calli indistinct.
impressions, puncture and texture of the intervals the same as with bactriana, only the first interval apically with fewer, lesser distinct and lesser regular transverse wrinkles. Meta-sternum Metasternum medio late excavatum (♂).
The first segment of the abdomen in the middle longwise broadly impressed, the apex of the last segment rectangularly truncated and triangularly impressed (♂). Pygidium Pygidium distincte arcuato-emarginatum. Pygidium distinctly arcuately emarginate.
hind tibiae curved, scarcely broadened at the end of the first third, without notches. Size Long. 8 mill.; lat. 2,6 mill.
Inhabits the town of Kjachta in the province of Transbaicalia in eastern Siberia (coll. Clavareau).

Appendix 2
Donacia fukiensis Goecke, 1944. Original description in German and translation into English.
Medium sized uniform dark bronzy shiny animals with an extremelyfinely pubescent pronotum, the males more slender and shorter than the females, which have a much lesser clubbed thickened femur and their 1 st abdominal segment is not flattened. The animals' habitus is very uniform, the formation of the single characters is very variable. Head Oberkiefer überragt die Oberlippe um etwas mehr als deren Länge, pechbraun, Kiefertaster gelb, bei einigen Stücken das letzte Glied an der Spitze braun.
Labrum ca. twice as broad as long, front margin slightly convexly rounded, basal margin with long setae, reaching beyond the front margin, front margin with shorter setae, in between without setae. Kopfschild vorn gerade, 2mal so breit wie die Seitenkante lang.
Head plate straight at front, twice as broad as the length of the side margin. The antennae are filiform, not very long, in males reaching farer than the middle of the elytra, in females they are significantly shorter. 2 nd segment the shortest, about half as long as the 1 st one, the 3 rd one about one fifth to one half longer than the 2 nd one, the 4 th one is one and a half times to double the length of the second one. The length relations of the single segments are quite variable to each other. Antennomeres yellow to dark brown. The 1 st to 6 th one with moderately dense hairs, the 7 th to 11 th one with more densely packed hairs. Die Fühlerhöcker sind abgeplattet, dazwischen befindet sich eine schmale Furche, die Abplattung ist mehr oder weniger glänzend, fast ohne Punkte oder mäßig dicht punktiert, dahinter befindet sich eine mehr oder weniger deutliche Vertiefung, die gegen die Fühlerhöcker durch eine querliegende Kante abgesetzt ist. Die Stirnhöcker sind ziemlich flach und breit. Äußere Gruben flacher oder tiefer, innere Gruben schwach entwickelt.
The antennal tubercles are flattened, with a narrow groove between them, the flattened part is more or less shiny, almost without punctures or moderately densely punctured, behind it there is a more or less distinct depression, which is separated against the antennal tubercles by a transverse ridge. The calli are quite flattened and broad. Outer grooves more flattened or deeper, inner grooves shallow. Stirn mäßig dicht punktiert und behaart, glänzend.
Eyes small, wide apart.
Broadest part of the pronotum at the anterolateral tubercles and approximately as broad as its length in the midst. However the pronotum of one specimen was considerably longer. Die Vorderecken sind gut entwickelt, sie ragen aber weder über den Vorderrand noch über die Seitenhöcker vor.
The anterior angles are well developed, but neither protruding beyond the anterior margin nor the lateral tubercles. Vorderrand leicht konvex, gegen die Scheibe nicht, oder durch eine feine, oft unregelmäßige Linie abgesetzt.
Anterior margin slightly convex, not distinctly separated against the disc, or by a subtle, often irregular line. Hinterecken mehr oder weniger gut entwickelt, wenig vorragend.
The disc of the pronotum is very variable, evenly shallowly domed, almost without a hint of a central groove or flattened and with a distinct longitudinal groove. The central groove neither reaches the anterior nor the posterior margin, at the most it peters out to a shallow or only indistinct impression ahead or rearmost. Anterior lateral tubercles distinct, against above slightly or scarcely, against the anterior angles distinctly, against backwards slightly separated. Posterior lateral tubercles poorly developed. Not densely, irregularly punctured, in between the punctures shiny. Often the punctures are more dense in the anterior and posterior part than in the middle part. But the density of the punctures is very different between single specimens. Der Halsschild ist behaart. Es befinden sich nämlich in den Punkten äußerst feine, kurze, sehr schwer sichtbare Borsten.
The pronotum is pubescent. For inside the punctures there are exceedingly delicate, short setae which are very difficult to be seen. Prothorax Die Episternen der Vorderbrust sind grob längs gerunzelt, der behaarte Fleck ist nur schwach behaart.
Elytra feebly domed from anterior to posterior, more distinctly and evenly towards the margins, twice as long as the breadth of both. Outer contour parallel from anterior to the second third, then evenly domed towards the singly rounded apices. Truncation indistinct.
The intervals are flattened and broad, shiny with flat, greatly separated transverse wrinkles and with very fine more or less dense micropuncture. The 1 st interval is almost glabrous with only weak transversal, longitudinal or diagonal wrinkles and margined on both sides with a ridge like a solid line in the last third.
The first impression at the suture is distinct only at some specimens, almost invisible at others. Other impressions are lacking besides the weakly developed humeral groove. Meta-thorax Die Unterseite der Hinterbrust ist beim ♂ herzförmig abgeplattet, beim ♀ gewölbt mit tiefer liegender Mittelfurche.
The 1 st abdominal segment is slightly longer at the ♂, at the ♀ longer by the half than the 2 nd to 5 th together, at the ♂ flattened and slightly impressed, at the ♀ domed. Das letzte Segment ist beim ♂ an der Hinterkante leicht eingedrückt, beim ♀ konvex vorgezogen ohne eigentliche Spitze.
The last segment is slightly impressed at the apical ridge at the ♂, at the ♀ convexly protruding without a distinctive peak. Die Unterseite des Hinterleibes ist glänzend, mäßig dicht punktiert und behaart.
The underpart of the abdomen is shiny, moderately densely punctured and pubescent. Das Pygidium ist abgestutzt und in der Mitte schwach ausgebuchtet.
The pygidium is truncated and slightly recessed in the middle.
The posterior femora are short, even at the ♂ they don't reach the apex of the elytra by far, anterior, middle and posterior femora much thickened like clubs especially at the ♂, at the ♀ more slender. Hinterschenkel mit einem kräftigen Zahn, der beim ♂breiter, beim ♀ schmaler und spitzer ist (siehe Abb. 5).
The 1 st and 3 rd tarsomere have about the same length, the 2 nd one is by a third shorter. Colour Die Tiere sind einheitlich dunkel bronzefarben, nur die Fühler gelb bis dunkelbraun, die Schienen und Tarsen und die Hinterschenkel von der Basis bis zur Mittel hellbraun.