“Paradoxal” New Genus and Species of the Family Nitidulidae (Polyphaga

ZOOSYSTEMATICA ROSSICA, 20(2): 274–298 25 DECEMBER 2011

“Paradoxal” new genus and species of the family Nitidulidae (Polyphaga: Coleoptera) from Afro-Madagascarean and Australian Regions “Парадоксальные” новые род и виды семейства Nitidulidae (Polyphaga: Coleoptera) из Афро-Мадагаскарских и Австралийских областей

A.G. KIREJTSHUK

А.Г. КИРЕЙЧУК

A.G. Kirejtshuk, Zoological Institute of the Russian Academy of Sciences, 1 Universitetskaya Emb., St Petersburg 199034, Russia; CNRS UMR 7205, Muséum National d’Histoire Naturelle, CP 50, Entomologie, 45 Rue Buffon, F-75005, Paris, France. E-mail: [email protected], [email protected]

In the paper described are two new species of the genus Amphicrossus Erichson, 1843 (A. microtuberculatus sp. nov. from Sudan and A. uhligi sp. nov. from Zambia: Amphicrossinae), one new species of the genus Meligethinus Grouvelle 1906 (M. zimbabwensis sp. nov. from Zimbabwe: Meligethinae), one new species from the genus Neopallodes Reitter, Reitter, 1884 (N. madagascarensis sp. nov. from Madagascar: Nitidulinae, Cyllodini) and one species of the new genus Gonoglypha gen. nov. (G. distinctissima sp. nov. from Australia (Queensland): Niti- dulinae, Cychramptodini). In the paper also some addition to the description of Meligethinus dolosus Grouvelle, 1919 from the eastern part of South Africa and some comments on sig- nificance of different characters, taxonomy and classifications of some groups of the family Nitidulidae are included. В статье описаны два новых вида из рода Amphicrossus Erichson, 1843 (A. microtuberculatus sp. nov. из Судана и A. uhligi sp. nov. из Замбии: Amphicrossinae), один новый вид рода Meligethinus Grouvelle 1906 (M. zimbabwensis sp. nov. из Зимбабве: Meligethinae), один новый вид рода Neopallodes Reitter, Reitter, 1884 (N. madagascarensis sp. nov. из Мада- гаскара: Nitidulinae, Cyllodini) и один новый вид из нового рода Gonoglypha gen. nov. (G. distinctissima sp. nov. из Австралии (Квинсленда): Nitidulinae, Cychramptodini). В статью так же включены дополнение к описанию Meligethinus dolosus Grouvelle, 1919 из восточной части Южной Африки и некоторые комментарии к значению различных при- знаков, таксономии и классификациям некоторых групп семейства Nitidulidae. Key words: Africa, Madagascar, Australia, Coleoptera, Nitidulidae, Amphicrossinae, Meli- gethinae, Nitidulinae, new genus, new species Ключевые слова: Африка, Мадагаскар, Австралия, Coleoptera, Nitidulidae, Amphicros- sinae, Meligethinae, Nitidulinae, новый род, новые виды

INTRODUCTION (Ecuador); Bolivitoxus Kirejtshuk, 2009 (Bolivia) and Microsoronia Kirejtshuk et The recently elaborated generic system Kurochkin, 2010 (Baltic amber); one ge- of the family (Kirejtshuk, 2008) was added nus of Cillaeinae: Brittonema Kirejtshuk, by some further contributions. There have 2011 (Australia); one genus of Epuraeinae: been published three genera of the subfam- Baltoraea Kurochkin et Kirejtshuk, 2010 ily Nitidulinae: Neohebascus Cline, 2009 (Baltic amber); one genus of Cybocephali-

© 2011 Zoological Institute, Russian Academy of Scienсes A.G. KIREJTSHUK. “PARADOXAL” NEW GENUS AND SPECIES OF NITIDULIDAE 275 nae: Pastillocenicus Kirejtshuk et Nel, 2008 During last dozens some undescribed (Oise amber); besides, three generic names genera and many new species, subfamily and were proposed in Meligethinae: Sebastian- generic attribution of which can be defined gethes Audisio, Kirk-Spriggs et Cline, 2008 due to some characters additional to the (South Africa); Restiopria Audisio, Jelínek traditional diagnoses of the taxa including et Cline in Audisio et al., 2011 (South Af- these new “paradoxal” forms have been re- rica) and Melipriopsis Kirejtshuk, 2011 covered. Nevertheless, relationship of these (Baltic amber), in press in addition to the new forms to the taxa where they are put classification published by Audisio et al. could be supposed with a rather high prob- (2009) with proposal of 22 new generic ability thanks to the additional characters names. In this classification of the last sub- considered in the paper. The present paper family all most taxa treated before as sub- deals with description of some such species genera were elevated to the generic level and one genus. The “paradoxal” forms pres- and some synonyms were reestablished as ent some difficulties for classification, al- valid genera. This large revision with many though they in addition to the fossil record important illustrations includes division can provide phylogenetic reconstruction “Generic (re) description and diagnosis” for with extremely valuable information. many taxa regarded by the authors as valid. Depositories. BMNH – The Natural However even these divisions have neither History Museum, London (formerly Brit- proper diagnosis nor comparison with clos- ish Museum of Natural History); MNHN – est taxa. As a result the concept of the au- Museum National d’Histoire Naturelle, thors of this classification remains unclear Paris; QM – Queensland Museum, Bris- and can be scarcely discussed. Each genus bane; ZIN – Zoological Institute of the Rus- included in it is provided with a short divi- sian Academy of Sciences, St Petersburg; sion “Phylogenetic position” that contains a ZMB – Zoologisches Museum an der Hum- description of placement of a certain taxon boldt Universität, Berlin. in the cladogram put in the end of this publication. In the legend to this cladogram the RESULTS authors wrote that it was based on “72 mor- phological characters”, but no list of these Family NITIDULIDAE Latreille, 1802 characters and their distribution among Subfamily AMPHICROSSINAE the taxa included are supplied. Fortunately, Kirejtshuk, 1986 each generic taxon is provided with a list of Genus Amphicrossus Erichson, 1843 the species included. In practice nobody can identify many of genera sensu Audisio and Type species: Nitidula ciliata Olivier, Cline (Audisio et al., 2009) without knowl- 1811, recent (after designation by C.T. Par- edge on the species attribution of specimens sons, 1943). under identification. Because of difficulties Notes. Two new species described below of use of this classification the author of this differ from all congeners in the character paper considers the system of all subfami- of sculpture on dorsal integument and re- lies as it was outlined in Kirejtshuk, 2008. duction of isolation of the median plate of At the same time almost all species men- their mesoventrite (this plate is important tioned by Audisio et al. (2009) are known diagnostic feature of Amphicrossus: Parsons, to the author of this paper and he takes into 1972 etc.). The lack of a clear isolation of consideration the concept of the mentioned the median plate of the mesoventrite co- authors and probable incongruence which incides with anterolateral continuations can appear if somebody tries to apply only very gently sloping in the anterior part and to one of these versions of the classification a more gentle curve of the median plane of of Meligethinae. the mesoventrite in general, while in the

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 274–298 276 A.G. KIREJTSHUK. “PARADOXAL” NEW GENUS AND SPECIES OF NITIDULIDAE congeners the median plate of the mesoven- Diagnosis. This new species differs from trite looks like a continuation of plane of all known congeners in the elongate and the metaventrite, which is sharply declined comparatively rather weakly convex body. anteriorly and separated from the anterior Besides, in contrast to almost all congeners parts of mesoventrite by distinct ridges. it has also the very short and shallow me- These features in addition to the not even- dian furrow in the anterior part of meso- ly convex elytra and comparatively weak ventrite between rather small paramedian convexity of their dorsum in general make plates. As the A. uhligi sp. nov. it has also the these species very distinct among the mem- not isolated median plate of mesoventrite, bers of the subfamily. Nevertheless differ- comparatively narrow and curved male ences in the structure of male genitalia and metafemur and particularly not punctured genital capsule as well as other peculiarities but microtuberculate dorsal integument. A. (see the diagnoses below) are evidence that microtuberculatus sp. nov. differs from the these new species are scarcely close relatives latter in the more slender body with the less and the similarity mentioned above did not arcuate sides, shallowly excised anterior appear as a sequence of a common ancestry. edge of pronotum, and subflattened dorsum (particularly pronotum), much finer micro- Amphicrossus microtuberculatus sp. nov. tuberculation of dorsal integument, outline (Figs 1–12) of anterior edge of labrum, subhemicircular Material examined. Holotype, male antennal club, presence of clear paramedi- (BMNH) – “Sudan, Nuba Mountains, Talodi, xii- an brushes at distal third of elytral suture 1921, Capt. F. Moysey”, “Brit. Mus., 1922–194”. in male, longer male pygidium, shape of

Figs 1–5. Amphicrossus microtuberculatus sp. nov. Holotype, male: apex of ultimate abdominal segment, ventral (1); anal sclerite, ventral plate and spiculum gastrale, ventral (2); tegmen, ventral (3); idem, lateral (4); idem, posterior (5). Scale bars: 1.4 mm (1), 0.7 mm (2–5).

Figs 6–12. Amphicrossus microtuberculatus sp. nov. Holotype, male: body, dorsal (6); idem, ventral (7); idem, lateral (8); sculpture of pronotum and elytra (9); head, dorsal (10); idem, ventral (11); prosternum and mesoventrite, ventral (12). Body length 7.4 mm.

Figs 13–19. Amphicrossus uhligi sp. nov. Holotype, male: body, dorsal (13); idem, ventral (14); idem, lateral (15); sculpture of pronotum and elytra (16); head, dorsal (17); idem, ventral (18); prosternum and mesoventrite, ventral (19). Body length 8.3 mm.

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 274–298 A.G. KIREJTSHUK. “PARADOXAL” NEW GENUS AND SPECIES OF NITIDULIDAE 279 ultimate labial and maxillary palpomeres, gidium ciliate by long, dense and compara- longer median plate and very short paratively thick hairs about as long as hairs on median plates, curved metafemora, longer underside. hypopygidium with only a very short and Head, pronotum and elytra without narrow shining median stripe along its puncturation, but with rather small and posterior edge and narrower movable lobe dense diffuse tubercles (protuberances), di- almost completely concealed under plate ameter of which is markedly less than half of the hypopygidium. By the general body of that in eye facets and which are becom- outline, presence of paramedian brushes ing inclined posteriorly on elytra, inter- at elytral suture in male, structure of ulti- spaces between them looking like about as mate visible male abdominal segment, male great as tubercles. Pygidium and exposed genital capsule and genitalia this new spe- abdominal tergite with very fine, shallow cies is similar and seems to be related to A. and dense punctures, interspaces between namibiensis Kirejtshuk, 1987 from Namibia them greater than a puncture diameter and but rather distinct from the latter in the very densely microreticulated. Underside lighter body, clear microtuberculation on with medium-sized, shallow, but distinct the dorsum (instead of usual puncturation punctures, interspaces between them 4–5 on the dorsum in both other species), much times as great as a puncture diameter and shorter dorsal pubescence, shallower exci- very smoothly microreticulated. sion of anterior edge of pronotum, lack of Head flattened and about as long as the adsutural lines on elytra and shape of lateral distance between moderately large eyes lobes of the tegmen. Analogous similarities (consisting of medium-sized facets), trans- and differences can be also observed com- verse diameter of the latter nearly as a third paring this new species and A. pubisetosus of the distance between eyes, its anterior Kirejtshuk, 1987 from Sudan, however, in edge transverse and with rounded lateral contrast to the new species, the latter also angles. Lobes of labrum slightly exposed, characterised by very distinct structure of subhemicircular and with a comparatively last abdominal segment with widely trans- deep excision between. Mandibles slightly verse apex of hypopygidium bearing a far exposed from under lobes of labrum. An- projecting median movable lobe. tennae somewhat shorter than head wide; Description of male (holotype). Length their club composing about 2/7 of total 7.4, breadth 2.6, height 1.2 mm. Feebly con- antennal length, suboval, slightly longer vex dorsally and moderately convex ven- than wide and with ultimate antennomere trally; dorsum light brownish to reddish; rather blunt at apex. Pronotum widest be- underside, appendages and abdominal ter- fore base and regularly very widely rounded gites nearly straw reddish; dorsum nearly anteriorly, with subflattened disk (to sub- mat and underside with a fat shine; dorsum depressed in the middle) and gently sloping with rather fine, suberect and slightly con- to narrowly bordered (not explanate) sides, spicuous yellowish hairs more than twice anterior edge weakly trapezium-like ex- as long as distance between their insertions cised, posterior edge shallowly emarginate, and, besides, there are sparse and markedly posterior angles widely rounded. Scutellum longer hairs diffuse on head and pronotum subpentagonal with rounded apex, almost but arranged in longitudinal rows on ely- two thirds as wide as long. Elytra nearly tra; underside with much longer, fine and 1 and 1/2 as long as wide combined, with moderately conspicuous hairs (particularly maximum width near midlength, distally on prosternum); elytra with paramedian gradually narrowing to obliquely truncate brushes at distal third of suture; sides of apices, which are forming a very shallow pronotum, elytra and abdominal segments sutural angle, steeply sloping to very nar- as well as apices of pygidium and hypopy- rowly subexplanate lateral edges, adsutural

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 274–298 280 A.G. KIREJTSHUK. “PARADOXAL” NEW GENUS AND SPECIES OF NITIDULIDAE lines not expressed. Pygidium slightly con- Genital capsule. Anal sclerite with arcu- vex, slightly wider than long and widely ate apex; ventral plate with very short lobes. subtruncate at apex. Aedeagus. Tegmen heavily sclerotised Ultimate labial palpomere about twice and rather long; penis membraneous and as long as thick, slightly and gradually without any clear armature in its inner sac. thickening apically. Ultimate maxillary pal- Etymology. The epithet of this new spe- pomere almost four times as long as thick, cies refers to a characteristic sculpture of its subcylindrical and slightly narrowing at dorsal integument. apex. Mentum subpentangular, somewhat more than twice as wide as long, rectilinear Amphicrossus uhligi sp. nov. lateral edges only slightly shorter than men- (Figs 13–25) tum at midline. Antennal grooves distinct and widely arcuate. Prosternum gently con- Material examined. Holotype, male (ZMB) – vex along the middle and with flat process “Zambia, 26–29.III.1993, 15°02´S/26°00´E, moderately widened at subtruncate apex, Chunga Camp, Kafue NP, lux, leg. U. Göllner”. which is about 5/6 as wide as antennal club. Diagnosis. This new species is rather Distance between mesocoxae somewhat similar to the previous one (see the above more than 2.5 times and that between meta- diagnosis). The aedeagus of it is more or coxae somewhat more than twice as great less similar to that in A. concolor Murray, as that between procoxae. Median plate of 1867 widely spread in the Afrotropics and mesoventrite somewhat longer than wide A. fuligorufus Kirejtshuk, 1995 from Zaire along its posterior edge and without ante- although lateral lobes of tegmen are rather rior ridges; paramedian plates of mesoven- different. Amphicrossus uhligi sp. nov. dif- trite rather short, about a third as long as fers from both in its more slender body with median plate and with a slightly traceable less convex dorsum and completely exposed short furrow between them. Metaventrite pygidium (not covered with the elytral subflattened along the middle, with subme- apices), characteristic sculpture of dorsal socoxal lines following closely the posterior integument, structure of mesoventrite and edge of cavity and slightly deviating only last abdominal segment (particularly in the at outer angle of metaventrite. Submeta- very short movable lobe at apex of hypopy- coxal line following closely posterior edge gidium); and also from the first in the some- of cavity. Abdominal ventrite 1 longest, al- what elongate antennal club and shape of most as long as ventrites 2 and 3 combined prosternal process; and from the second in and somewhat longer than hypopygidium, the subunicolourous body. which is subtruncate at apex and bearing a Description of male (holotype). Length very small smooth stripe along the middle 8.3, breadth 3.5, height 1.8 mm. Feebly con- of its posterior edge. Movable lobe of hypo- vex dorsally and moderately convex ven- pygidium comparatively small and narrow, trally; dorsum light brownish to reddish; scarcely exposed from under its apex. Epi- underside, appendages and abdominal ter- pleura slightly elevated laterally and about gites nearly straw reddish; dorsum nearly as wide as antennal club at base. mat and underside slightly shining; dorsum Tibiae of usual structure, about as wide with rather fine, subrecumbent and slightly as antennal club at apex, comparatively nar- conspicuous yellowish hairs about twice as row and rather long; spurs rather strong and long as distance between their insertions stout. Femora of usual shape and compara- and, besides, there are sparse and markedly tively narrow, 1.7–2.2 times as wide as tibi- longer hairs diffuse on head and pronotum ae. Protarsus about half as wide as protibia, but arranged in longitudinal rows on elytra; meso- and metatarsi somewhat narrower; underside with much longer, fine and mod- claws simple and about as long as tibial spurs. erately conspicuous hairs; elytra without

Figs 20–25. Amphicrossus uhligi sp. nov. Holotype, male: anal sclerite, ventral plate and spiculum gastrale, ventral (20); tegmen, ventral (21); idem, lateral (22); idem, posterior (23); penis trunk, dorsal (24); armature of inner sac of penis (25). Scale bar: 0.7 mm. paramedian brushes at distal third of suture; and cellularly microreticulated interspaces. sides of pronotum, elytra and abdominal Elytra with tubercles shallower and sparser segments as well as apices of pygidium and than those on head and pronotum, becoming hypopygidium ciliate by long, dense and inclined posteriorly and partly dislodged by comparatively thick hairs about as long as very shallow punctures along sides and at hairs on underside. apices, interspaces between them somewhat Head and pronotum and elytra without greater than tubercles and with fine cellular puncturation, but with small and dense dif- microreticulation. Pygidium and exposed fuse tubercles (protuberances), diameter of abdominal tergite with very fine and dis- which is nearly as great as that in eye fac- tinct punctures, interspaces between them ets, interspaces between them looking like about three times as great as a puncture di- about as great as tubercles. However along ameter and very densely microreticulated. sides and anterior edge of frons there are Underside with punctures somewhat larger visible clear small punctures with finely than those on pygidium, interspaces be-

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 274–298 282 A.G. KIREJTSHUK. “PARADOXAL” NEW GENUS AND SPECIES OF NITIDULIDAE tween them 4–5 times as great as a puncture the middle and with flat process moderately diameter, smooth to very smoothly micro- widened at widely rounded to subtruncate reticulated on thoracic sclerites, and finely apex, which is about as wide as antennal and cellularly reticulated on abdomen. club. Distance between mesocoxae about Head flattened and about as long as the three times and that between metacoxae distance between moderately large eyes about 1 and 1/2 as great as that between (consisting of medium-sized facets), trans- procoxae. Median plate of mesoventrite verse diameter of the latter nearly as a third markedly shorter than wide along its poste- of the distance between eyes, its anterior rior edge and without anterior ridges; para- edge transverse and with rounded lateral median plates of mesoventrite short, about angle. Lobes of labrum with inclined lateral half as long as median plate and with a clear sides and a comparatively narrow excision furrow between them. Metaventrite subflat- between. Mandibles slightly exposed from tened along the middle, with very subme- under lobes of labrum. Antennae somewhat socoxal line following closely the posterior shorter than head wide; their club com- edge of cavity and slightly deviating only at posing about 2/7 of total antennal length, outer angle of metaventrite. Submetacoxal subovoid, about 1 and 1/3 as long as wide line following closely the posterior edge of and with ultimate antennomere angular at cavity. Abdominal ventrite 1 longest, almost apex. Pronotum widest before base and reg- as long as ventrites 2 and 3 combined and ularly very widely rounded anteriorly, with nearly twice longer than hypopygidium, slightly convex disk and gently sloping to which is subtruncate at apex and bearing a narrowly bordered (not explanate) sides, moderately small smooth stripe along the anterior edge moderately trapezium-likely middle of its posterior edge. Movable lobe excised, posterior edge shallowly emargin- of hypopygidium comparatively small and ate, posterior angles widely rounded. Scu- rather wide, slightly exposed from under its tellum subpentagonal with rounded apex, apex. Epipleura slightly elevated laterally more than twice as wide as long. Elytra and about 1 and 1/3 as wide as antennal nearly 1 and 3/8 as long as wide combined, club at base. with maximum width near midlength, dis- Tibiae of usual structure, somewhat tally gradually narrowing to very widely wider than antennal club at apex, compara- and separately rounded apices, which are tively narrow and rather long; spurs rather forming a moderately deep sutural angle, strong and stout. Femora of usual shape steeply sloping to very narrowly subex- and comparatively narrow, less than twice planate lateral edges, adsutural lines slight- as wide as tibiae. Protarsus only slightly ly traceable only at distal end. Pygidium narrower than protibia (about as wide as slightly convex, markedly wider than long antennal club), meso- and metatarsi some- and widely subtruncate at apex. Transverse what narrower; claws simple and about as apex of anal sclerite scarcely exposed from long as tibial spurs. under pygidial apex. Genital capsule. Anal sclerite with trun- Ultimate labial palpomere about twice cate apex; ventral plate with four moder- as long as thick, subcylindrical. Ultimate ately raised lobes. maxillary palpomere almost three times as Aedeagus. Tegmen heavily sclerotised long as thick, subcylindrical and scarcely and rather long; penis membraneous and narrowing only at apex. Mentum subpen- with two slightly sclerotised stripes in ar- tangular, somewhat more than twice as mature of its inner sac. wide as long, subrectilinear lateral edges Etymology. The epithet of this new species only slightly shorter than mentum at mid- is devoted to Manfred Uhlig, an old friend of line. Antennal grooves distinct and widely mine who collected many interesting beetles arcuate. Prosternum gently convex along in Africa of the South Hemisphere.

Subfamily MELIGETHINAE sp. nov. is more similar to some species of C.G. Thomson, 1859 Cyclogethes rather than to species of the Genus Meligethinus Grouvelle, 1906 rest genera. Meligethinus zimbabwensis sp. nov. is characterised by many characters Type species: Meligethinus humeralis similar to those in species of Meligethinus Grouvelle, 1906, recent (by monotypy). widely spread mainly in the Palaeotropics Notes. The new species of this genus and Micropria Grouvelle, 1899 from the having a peculiar shape of the (very wide Afrotropical Region, but differs from the bi-arcuate) depressions at the base of second in the somewhat larger body, striate pygidium shares also some other pecu- elytra, bi-arcuate depressions at the base of liarities with some representatives of the pygidium, longer frons and narrower pros- genera from the Indo-Malayan Region ternal process. The genus Meligethinus is (Cryptarchopria Jelínek, 1975; Cyclogethes rather diverse, but the new species differs Kirejtshuk, 1979; Horakia Jelínek, 2000; from all congeners in the bi-arcuate depres- Kabakovia Kirejtshuk, 1979), particularly sions at the base of pygidium and from most similar in the outline of dorsal sclerites, of them in the striate elytra. Nevertheless, transversely striate elytra, characteris- a trend to appearance of a transrugosity tic prosternal process and mesoventrite, is observed in some African species of the (sub) parallelepiped-shaped meso- and genus, although the completely striate in- metatibiae, acute apex of penis trunk and tegument of elytra was found only in one deeply excised tegmen. Nevertheless it dif- known species, Meligethinus dolosus Grou- fers from Cryptarchopria, Cyclogethes and velle, 1919 from the eastern part of South Kabakovia as from all other genera of the Africa or also south of Zimbabwe, which subfamily in the pygidium with a pair of has the simple arcuate depressions at base the above mentioned depressions at base, of its pygidium, as in most congeners. The and also from Cryptarchopria in the lack of differences between the latter and new prosternal median plate and unexpressed species are defined in the below diagnosis. sexual dimorphism in antennae; and also All the considered genera could be related from Kabakovia in the pair of the simply ar- to the widespread Meligethinus, species of cuate depressions at base of hypopygidium which are distributed as in many areas of (but not bi-arcuate depressions). Besides, Africa, Mediterranean, Palaearchearctic in contrast to Meligethinus, Cyclogethes has (East Chinese) Province and Indo-Malay- the dorsal integument between punctures an Region. The species of Meligethinus, Ka- very smooth and shining; and also Horakia bakovia, and Cryptarchopria are associated has not a pair of arcuate depressions at base with palm inflorescences (Audisio, 1993; of hypopygidium (structure of base of py- Kirejtshuk & Kabakov, 1997) and Hora- gidium is unknown in Horakia). Some spe- kia presumably also (Jelínek, 2000), while cies of the subgenus Clypeogethes Sholtz, there is nothing known on host plant of Mi- 1932 of the genus Meligethes Stephens, cropria and Cyclogethes. In the cladogram 1832 sensu Kirejtshuk, 1992 (i.e. includ- of Audisio et al. (2009) Micropria and Cy- ing some “genera” by Audisio & Cline in clogethes were placed in the clade different Audisio et al., 2009) have the same differ- from that uniting Kabakovia, Horakia and ences from the species of Meligethinus zim- Cryptarchopria together with Meligethinus, babwensis sp. nov. as those of Cyclogethes, however the argumentation for the separa- and species of Clypeogethes frequently tion of these clades has not been published. differ also in the shape of body, armature Besides, these cladogram and “phyloge- of protibiae, emarginate or excised ante- netic” interpretation proposed by Audisio rior edge of frons and other characters. et al. greatly conflict with the fossil record The aedeagus of Meligethinus zimbabwensis (Kirejtshuk, 2011 in press).

Meligethinus zimbabwensis sp. nov. regular, somewhat obliquely transverse (Figs 26–32, 37–40) striae and extremely small punctures disposed behind them, interspaces between Material examined. Holotype, male (ZMB) – striae about two diameters of eye facets and “Zimbabwe: Victoria falls, Zambesi Nat. finely alutaceous, striae becoming gradu- Park, Camp, 11–13.XII.1993, 700 m, ü.N., ally denser towards apices. Pygidium and 17°53´S/25°49´S, J. Deckert, lux”; 1 para- abdominal ventrites with small, shallow, type, female (ZIN) – “Zimbabwe: 12.xii.1993, but distinct punctures, interspaces between 17°53´S/25°49´S, Victoria Falls, Zambesi-NP- them greater than a puncture diameter and Camp, lux, M. Uhlig”. very smoothly microreticulated. Proster- Diagnosis. This new species differs from num, metaventrite and abdominal ventrite another congener with the transversely 1 with extremely fine puncturation and striate elytra (M. dolosus) in the dark ely- more or less smooth. tra with somewhat sparser striae, somewhat Head subflattened and about as long longer head with somewhat larger eyes, as the distance between moderately large shorter antennae with longer antennomere eyes (consisting of rather fine facets), 4, shorter antennal club, longer lateral edge transverse diameter of the latter somewhat of mentum, longer ultimate maxillary pal- less than half of distance between eyes, its pomere, markedly weaker postocular de- anterior edge finely bordered, transverse pressions, shape of prosternal process, con- and with rounded lateral angles. Lobes of tour of submesocoxal line, narrower tibiae, labrum and mandibles slightly exposed deeply excised apex of male hypopygidium, from under lobes of labrum. Antennae structure of the male genitalia and also somewhat shorter than 3/4 of head width, slightly more conspicuous pubescence and antennomere 3 markedly shorter than an- slightly finer puncturation (particularly on tennomere 4, the latter about as long as the prosternum, metaventrite and abdomi- antennomere 1–3 combined; their club nal ventrite 1), which do not show distinct composing about 1/4 of total antennal transverse rows. length, subovoid, about 1 and 1/3 as long Description of male (holotype). Length as wide and with ultimate antennomere 2.7, breadth 1.4, height 0.7 mm. Moderately transversely truncate. Pronotum widest convex dorsally and moderately convex at base and regularly rounded anteriorly, ventrally; light brownish to reddish; scu- with evenly vaulted disk and gently slop- tellum and elytra dark brown to blackish ing to narrowly subexplanate sides, an- with an adsutural oval light spot at the apex terior edge moderately trapezium-likely of each elytron; appendages light reddish excised, posterior edge truncate with a (straw), but antennal club dark; integu- shallow sinuation at each posterior angle, ment with a fat shine and covered with ex- which is slightly projecting posteriorly. tremely fine greyish hairs 3–4 times as long Scutellum subtriangular with rounded as distance between their insertions and on apex, almost twice as wide as long. Elytra elytra their insertions located behind trans- about 1 and 1/5 as long as wide combined, verse striae. with maximum width in anterior third, Head and pronotum with rather small distally gradually narrowing to obliquely and distinct punctures much smaller than truncate apices, which are forming a very eye facets in diameter, interspaces between shallow sutural angle, steeply sloping to them on head about half as great as a punc- very narrowly subexplanate lateral edges, ture diameter and on pronotum somewhat adsutural lines distinct at distal 2/5. Py- greater, smoothly alutaceous; on head and gidium slightly convex, widely rounded at pronotum there are scarcely observed clear apex. Arcuate apex of anal sclerite scarcely transverse rows of punctures. Elytra with exposed from under pygidial apex.

Figs 26–32. Meligethinus zimbabwensis sp. nov. Holotype, male: prosternal process, ventral (26); mentum, ventral (27); pygidium, dorsal (28); tegmen and penis trunk, lateral (29); tegmen, ventral (30); penis trunk, dorsal (31); paratype, female: ovipositor, ventral (32). Scale bars: 1.0 mm (26), 0.5 mm (27–32).

Ultimate labial palpomere almost four inner edge wide, somewhat S-shaped, post- times as long as thick. Mentum subpen- ocular depression moderately deep. Pro- tangular, more than twice as wide as long, sternum gently convex along the middle lateral edges about 2/3 as long as mentum and with process slightly widened at suban- at midline. Antennal grooves distinct along gular apex, its maximum width somewhat

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 274–298 286 A.G. KIREJTSHUK. “PARADOXAL” NEW GENUS AND SPECIES OF NITIDULIDAE less than width of antennal club. Distance 5.XII.1995, KwaZulu-Natal, Hluhluwe Game between mesocoxae about 1.5 times that be- Res., 28°2´S/32°05´S, F. Koch”. tween metacoxae about 2.5 times as great as Diagnosis. See the diagnosis of the previ- that between procoxae. Metaventrite sub- ous species. flattened and with a weak and narrow medi- Addition to description of male. Length an depression, with very distinct submeso- 2.4–2.8, breadth 1.3–1.2, height 0.6 mm. coxal line following close the posterior edge Moderately convex dorsally and moderate- of cavity, subrectilinerly deviating only at ly convex ventrally; subunicolour yellowish outer angle of metaventrite and reaching to reddish; appendages straw yellowish to the middle of inner edge of metepisternum. light reddish; integument with a fat shine Submetacoxal line following closely poste- and covered with extremely fine greyish rior edge of cavity. Abdominal ventrite 1 al- hairs 1.5–2.0 times as long as distance be- most as long as ventrites 2–4 combined and tween their insertions and on elytra their nearly twice longer than hypopygidium, insertions located behind transverse striae. which is deeply excised at apex. Epipleura Head and pronotum with moderately gradually narrowing distally, subhorizontal small and distinct punctures, somewhat and almost 1.5 times as wide as antennal smaller than eye facets in diameter, inter- club at base. spaces between them on head about half as Tibiae about as wide as antennal club great as a puncture diameter and on pro- and moderately long; protibia finely crenu- notum somewhat greater, smoothly aluta- late along outer edge, meso- and metatibiae ceous; on head and particularly on prono- with a row of small very short and dense tum there are observed slight and irregular hairs along outer edge, spurs comparatively transverse rows of punctures. Elytra with small and stout. Femora of usual outline regular somewhat obliquely transverse stri- and rather wide, 2.5–3.0 times as wide as ae and extremely small punctures disposed prosternal process. Tarsi rather long (about behind them, interspaces between striae 2/3 as long as tibiae), tarsomeres 1–3 mod- about 1.5 diameters of eye facets and finely erately lobed, claw simple and narrow; pro- alutaceous, striae becoming gradually dens- tarsus about 2/3 as wide as protibia, meso- er towards apices. Pygidium and abdomi- and metatarsi somewhat narrower. nal ventrites with small, shallow, but dis- Aedeagus. Tegmen and penis trunk well tinct punctures, interspaces between them sclerotised and very long. greater than a puncture diameter and very Female. Length 2.6 mm. Externally dif- smoothly microreticulated. Prosternum, fers from the male only in the lack of the meta ventrite and abdominal ventrite 1 with median depression on metaventrite, a small fine puncturation and smoothly alutaceous. median excision at the apex of pygidium Head subflattened and about 4/5 as long and widely rounded apex of hypopygidium. as the distance between moderately large Ovipositor moderately sclerotised. eyes (consisting of rather fine facets). An- Etymology. The epithet of this new spe- tennae about as long as head wide, antenno- cies is formed from the name of country of mere 3 markedly longer than antennomere its origin. 4, the latter about as long as antennomere 1–3 combined; their club composing about Meligethinus dolosus Grouvelle, 1919 2/7 of total antennal length, suboval, about (Figs 33–36, 41–43) 1 and 1/5 as long as wide and with ultimate antennomere transversely truncate. Elytra Material examined. Paralectotype, male (MNHN) – “Kentani, H.P. Abornelly”, “dolo- about 1 and 1/3 as long as wide combined, sus Grouv.” (lectotype was designated in the adsutural lines distinct at distal 1/3. Ulti- collection of South African Museum by Coo- mate labial palpomere almost twice as long per, 1980); 1 male (ZMB) – “SÜDAFRIKA: as thick. Mentum subpentangular, more

Figs 33–36. Meligethinus dolosus. Specimen from ZMB, male: mentum, ventral (33); prosternal process, ventral (34); tegmen, ventral (35); penis trunk, dorsal (36). Scale bar: 0.5 mm. than twice as wide as long, lateral edge Aedeagus. Tegmen and penis trunk well about half as long as mentum at midline. sclerotised and very long. Prosternum gently convex along the middle and with process rather widened at subtrun- Subfamily NITIDULINAE Latreille, 1802 cate apex, its maximum width somewhat Tribe CYLLODINI Everts, 1898 more than width of antennal club. Distance Genus Neopallodes Reitter, 1884 between mesocoxae about twice that between metacoxae about four times as great Type species: Pallodes hilleri Reitter, as that between procoxae. Metaventrite 1877, recent (after designation by Kirejt- with a wide median depression and distinct shuk, 1994) submesocoxal line following close the posterior edge of cavity, arcuately deviating only Neopallodes madagascarensis sp. nov. at outer angle of metaventrite and reaching (Figs 44–59) the middle of inner edge of metepisternum. Submetacoxal line following closely poste- Material. Holotype, female (MNHN) – “Par- rior edge of cavity. Hypopygidium bisinuate is Museum, Madagascar, Goudot, 1834”, “Pal- at apex. Tibiae clearly wider than antennal lodes sp. nr.” (handwritten by A. Grouvelle). club and moderately long. Femora of usual Diagnosis. This new species is very dis- outline and rather wide, about 2.5 times as tinct among all congeners due to its widely wide as prosternal process. Tarsi moderate- rounded posterior angles of pronotum, com- ly long (about 3/4 as long as tibiae), male paratively long antennae with club includ- protarsus about 2/3 as wide as protibia. ing six antennomeres (while the rest conge-

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 274–298 288 A.G. KIREJTSHUK. “PARADOXAL” NEW GENUS AND SPECIES OF NITIDULIDAE ners have only 3–4 segments) and compos- na with very fine and very sparse punctures, ing about 2/3 of total antennal length, and with interspaces between them completely also due to the extremely short valvifer of smooth. Pygidium and abdominal ventrites the ovipositor. Besides, the body size of this with very small and very sparse punctures, new species is smallest among the conge- finely alutaceous. ners known from Madagascar (Kirejtshuk, Head subflattened, without a trace of 2008) – the rest members of the genus are at frontoclypeal suture, somewhat longer than least as long as 3.0 mm. The most Madagas- distance between eyes. Labrum as a trans- car species of the genus Neopallodes have the verse stripe about four times as wide as long, trapezoid scutellum with transverse apex, with rounded lateral angles and a narrow and only N. incertus (Grouvelle, 1906); N. median fissure. Antennae about 1.5 times as niger (Grouvelle, 1906); N. nitidulus (Grou- long as head wide; their 6-segmented club velle, 1906) and N. variabilis (Grouvelle, composing 2/3 of total antennal length. 1896) (although the last species is known Pronotum gently and arcuately narrowed only from Nossi-Bé) have the subtriangu- to apex, its anterior angles weakly project- lar scutellum. Nevertheless, Neopallodes ing and posterior ones widely rounded, madagascarensis sp. nov., in addition to the base viewed as a median projection some- differences mentioned above, differs from what covering scutellar base. Scutellum them in the darker subunicolourous body, subtriangular with widely rounded apex. and also differs from the first in the pres- Elytra widely rounded at sides, obliquely ence of small punctures between longitudi- and widely rounded apices, forming as clear nal rows of punctures on elytra and in the sutural angle; adsutural lines very short middle of metaventrite as well as in the lon- at only apices. Pygidium very convex and ger metatarsus; from the second in the out- rounded at apex. line of labral lobes and narrow femora; from Ultimate labial palpomere slightly nar- the third in the dorsal puncturation, which rowing apically, about three times as long is not reduced; from the fourth in the very as thick. Antennal grooves sharply outlined sharp prosternal carina along the middle, and gently convergent posteriorly. Men- longer ultimate labial palpomere and pres- tum trapezoid, widely rounded at sides and ence of puncturation on thoracic sterna. anterior edge, widest at base, 1 and 1/3 as Description of holotype (female). Length wide as long. Prosternum with distinct 2.8, breadth 1.8, height 1.2 mm. Body rather carina reaching anterior edge of proster- convex dorsally and moderately ventrally; num, its process strongly widened before unicolourous dark chestnut brown with subtruncate to widely rounded apex. Dis- slightly lighter appendages; rather shining; tance between mesocoxae about twice and dorsum glabrous; underside with very sparse, that between metacoxae about 2.5 times as very long and nearly inconspicuous hairs. great as that between procoxae. Mesoven- Head and pronotal surface with very trite “roof”-like, with an elevated smooth small, very sparse and moderately shallow stripe rather than carina in posterior half. punctures, much smaller than eye facets, al- Metaventrite gently and evenly convex, though some punctures larger than the rest its posterior edge shallowly emarginate. (nearly reaching eye facets in diameter), Submesocoxal line follow the posterior edge interspaces between them very finely alu- of mesocoxal cavity; intercoxal line arcuate taceous. Elytral surface with regular simple and away from anterior edge of metaven- longitudinal rows of distinct punctures trite on distance comparable with distance (slightly smaller than eye facets in diam- between mesocoxae. Abdominal ventrite 1 eter), between them very small punctures about as long as hypopygidium and nearly are dispersed, intervals between rows finely twice as long as each of ventrites 2–4. Hy- alutaceous to almost smooth. Thoracic ster- popygidium widely rounded at apex. Epi-

Figs 37–43. Meligethinus zimbawensis sp. nov., holotype, male: body, ventral (37); paratype: idem, dorsal (38); anterior part of frons, dorsal (39); holotype: hypopygidium (40). Body length holotype 2.7 and paratype – 2.6 mm. Meligethinus dolosus, paralectotype (MNHN), male: body, dorsal (41); paratype: idem, ventral (42); anterior part of frons, dorsal (43). Body length 2.8 mm.

Figs 44–50. Neopallodes madagascarensis sp. nov., holotype, female: body, dorsal (44); side of pronotum and elytron, dorsolateral (45); prosternum and head, ventral (46); metaventrite and interme- diate leg, ventral (47); abdomen, ventral (48); posterior leg, ventral (49); ovipositor, ventral (50). Body length 2.8 mm; ovipositor length 0.6 mm.

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 274–298 A.G. KIREJTSHUK. “PARADOXAL” NEW GENUS AND SPECIES OF NITIDULIDAE 291 pleura slightly narrower than antennal club laterally and narrowly subexplanate at lat- at base, strongly sloping downwards. eral edges, its anterior angles rounded and Protibia straight and without promi- slightly projecting, posterior angles widely nent subapical tooth (about half as wide as rounded; its anterior edge gently convex antennal club), mesotibia somewhat wider to shallowly bi-emarginate; posterior edge and subtriangular, metatibia nearly as wide somewhat convex; anterior and lateral edg- as protibia, straight, with subtruncate apex. es extremely narrowly bordered and ante- Femora of usual shape, pro- and metafemora rior one looking like a rather thin fold cov- about 2.5 times, mesofemur about twice as ering elytral base. Elytra incomplete, with wide as corresponding tibiae. Metatarsus arcuately subtruncate apices remaining the about as long as metatibia. most part of pygidium uncovered, with nar- Ovipositor moderately sclerotised. rowly explanate and bordered sides. Pygid- ium slightly convex and widely rounded to Tribe CYCHRAMPTODINI subtruncate at apex. Prosternum strongly Kirejtshuk et Lawrence, 1992 carinate along the middle and this carina Genus Gonoglypha gen. nov. continuing on very narrow process, which is not curved along procoxae and slightly Type species: Gonoglypha distinctissima widened at subacute apex; prohypomera sp. nov. strongly slopped laterally. All pairs of coxae Diagnosis. Body comparatively medium- narrowly separated. Mesoventrite subflat- sized (4.8 mm), subellipsoid, strongly con- tened in posterior half. Metaventrite some- vex dorsally and slightly convex ventrally. what elevated along the middle and with Dorsum rather finely and very sparsely large semicircular depressions before meta- punctured, alutaceous or finely microretic- coxae, with a distinct median suture (discri- ulated; elytral puncturation not arranged men) along the whole length and angularly in longitudinal rows; underside finely and excised posterior edge between metacoxae. sparsely punctured, smoothed between Intercoxal line between mesocoxae un- punctures; body very shortly pubescent and raised. Epipleura subhorizontal. Hypopy- with ciliate pronotal and elytral sides. Head gidium very widely rounded at apex. Legs slightly convex to subflattened and with rather short and wide. Tibiae very flat and medium-sized eyes, slightly narrowed at rather wide, with rather stout spurs but base, slightly projecting anteriorly, lateral without teeth and setae along outer edge. lobes of frons widely covering antennal in- Trochanters very transverse. Femora very sertions; anterior edge of frons looking like wide and subflattened. Tarsi comparatively smooth shining stripe deeply excised in the short, with widely lobed tarsomeres 1–3 middle; antennal grooves deep and sharply and simple claws. Tegmen as slightly curved outlined, their inner ridge extended later- plate without apical excision; penis trunk ally and partly cover flagellomeres; mentum rather short and truncate at apex. transverse, widely rounded at sides and ex- Comparison. The combination of the cised at anterior edge; pregenal process at characters of this new genus makes it ex- hypostomal sinus moderately wide. Labrum tremely isolated among the members of the not exposed. Ultimate labial and ultimate subfamily Nitidulinae. Particularly, absence maxillary palpomeres rather long and sub- of exposed labrum reminds such peculiarity cylindrical. Antennae 11-segmented, bear- in the subfamily Cryptarchinae Thomson, ing 3-segmented compact and oval club, 1859. At the same time the narrow carinate without any evident trace of sexual dimor- prosternal process, flattened (not carinate or phism. Pronotum subtrapezoid from above medially swollen) mesoventrite, structure and with subhemicylindrical surface, slight- labium and underside of head, medially ele- ly narrowing anteriorly, steeply sloping vated metaventrite with lateral depressions

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 274–298 292 A.G. KIREJTSHUK. “PARADOXAL” NEW GENUS AND SPECIES OF NITIDULIDAE for movement of metafemora, and especially extended inner ridges, comparatively long the structure of legs (very transverse tro- prosternal process and subhorizontal epi- chanters and strongly flattened tibiae with- pleura. Besides, Gonoglypha gen. nov. dif- out any armature along their outer edge), fers from Cylindroramus Kirejtshuk et Law- widely rounded posterior angles of prono- rence, 1992 in the more robust (not narrow- tum in this new genus are more similar to ly cylindrical) body with prohypomera not members of the endemic Australian tribe strongly vertical and not strongly elevated Cychramptodini of the subfamily Nitiduli- posteromesal portion of metaventrite; from nae and taking together have no parallels in Miskoramus Kirejtshuk et Lawrence, 1992 other groups of sap beetles, although some in the more slender and subellipsoid body, of these features can be separately observed subcylindrical elytra without adsutural in other groups of the family. The members lines and narrow prosternal process; from of this tribe are similar to members of the Cychramptodes Reitter, 1874 in the 11-seg- Cyllodes complex (Cyllodini) in that both mented antennae, subellipsoid body, subcy- groups have the prohypomera and elytral lindrical elytra with diffuse puncturation epipleura strongly and sharply declined and and without adsutural lines, not strongly ventrally produced (except for the new ge- elevated posteromesal portion of metaven- nus with subhorizontal epipleura), so that trite and narrower femora. the legs may be completely concealed be- The new genus shares many similarities neath the body; also these two groups have in the body shape and some organs with the reduced dorsal vestiture, distinct or indis- cyllodine genera Amborotubus Leschen et tinct rows of elytral punctures, enlarged an- Carlton, 2004 from Bolivia and Cerylollodes tenna1 club, and a tendency for the coxae Kirejtshuk, 2006 from New Zealand. Nev- to be approximate. The strongly declined ertheless it differs from both in the carinate and almost opisthognathous head of some prosternum with continuation of the medi- Cychramptodini resembles the condition an carina to apex of the prosternal process, in the Cyllodes complex (e.g. Viettherchnus flattened (not carinate) mesoventrite and Kirejtshuk, 1985 from the Indo-Malayan somewhat medially elevated metaventrite Region) and in the cryptarchine genus with lateral depressions for movement of Arhinella Kirejtshuk, 1981 from Equato- metafemora (important differences between rial Africa (Kirejtshuk & Lawrence, 1992). Cyllodini and Cychramptodini), as well as The structure of aedeagus of Gonoglypha in the much more convex body, characteris- distinctissima gen. et sp. nov., particularly tic structure of epicranium and its append- the short penis trunk and setae along the ages (outline of frons, eyes, very deep anten- truncate apex of tegmen, support this con- nal grooves, labial palpi and not exposed la- cept. However, the presence of pubescence brum), presence of dorsal pubescence, ciliate on dorsal integument ciliate pronotal and pronotal and elytral sides and subhorizontal elytral sides, subhorizontal elytral epipleu- (not elevated laterally) epipleura, although ra and lack of intercoxal line between me- metaventrite of Amborotubus and Cerylol- socoxae (characteristic of both nitiduline lodes have some small depressions at pos- tribes considered) in the new genus present terior angles of metaventrite. Gonoglypha essential problems for this interpretation. gen. nov. also differs from Amborotubus in This new genus differs from all three the trapezoid tibiae, lobed tarsi and lack of genera of Cychramptodini in the head with adsutural lines on elytra; from Cerylollodes lateral lobes of frons very far laterally ex- in the subhypognathous position of head, tended over antennal insertions, a deep very wide femora, very wide and very flat median excision of its anterior edge, pres- tibiae without setae along their outer edge, ence of dorsal pubescence, very deep anten- diffuse puncturation on elytra and very pe- nal grooves partly covered with laterally culiar structure of aedeagus.

Etymology. The name of the new genus is convex dorsally and moderately ventrally; formed from the Greek “gonos” (offspring) – subunicolourous light reddish with dark- “glypha” (engraving); gender feminine. ened (almost dark brown) apical parts of elytra; integument with a slight fat lustre; Gonoglypha distinctissima sp. nov. dorsum with very fine and nearly incon- (Figs 55–57, 60–66) spicuous long semi-erect hairs, about 3–5 times as long as distance between their in- Material. Holotype, male (QM) – “Mossman Bluff Track, 5–10 km W. Mossmann, N. Qld, 20 sertions, although pygidium with very fine, Dec 1989 – 15 Jan 1990, Monteith, Thompson & dense and very short hairs; pronotal and Anzses, site 9, 1260 m, fit. intercept”. elytral sides ciliate with somewhat thicker Description of holotype (male). Length and denser hairs; underside markedly lon- 4.8, breadth 2.0, height 1.2 mm. Body rather ger suberect hairs, which are rather thin

Figs 51–59. Neopallodes madagascarensis sp. nov., holotype, female: right antenna, ventral (51); prosternal process, ventral (52); mentum and labium, ventral (53); protibia, ventral (54); Gonog- lypha distinctissima gen. et sp. nov., holotype, male: head, dorsal (55); mentum and labium, ventral (56); prosternal process, ventral (57); Amborotubus clarkei, holotype, female (Arthropod Museum, Louisiana State University, Baton Rouge): head, dorsal (58); right elytron, lateral (59). Scale bar: 0.9 mm (51–55, 58, 59), 0.4 mm (56, 57).

Figs 60–66. Gonoglypha distinctissima gen. et sp. nov. Holotype, male: body, dorsal (60); idem, lateral (61); idem, ventral (62); head, dorsal (63); idem, ventral (64); tegmen and penis trunk, ventral (65); thorax, ventral (66). Body length 4.8 mm; tegmen and penis trunk length 0.4 mm.

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 274–298 A.G. KIREJTSHUK. “PARADOXAL” NEW GENUS AND SPECIES OF NITIDULIDAE 295 and slightly conspicuous in the middle of illary palpomere about four times as long thoracic sclerites, but thicker and more as thick. Ventral tentorial fossae and gular conspicuous on metepisterna and abdomi- sutures rather distinct. Inner ridge of an- nal ventrites. tennal grooves extended laterally nearly Head and pronotal surface with very regularly arcuate; antennal grooves com- small, rather sparse and moderately shallow pletely received flagella. Prosternal pro- punctures, much smaller than eye facets, in- cess somewhat extended behind the pos- terspaces between them 2–5 times as great terior edges of procoxae, procoxal cavities as eye facets in diameter (denser on head looking like incompletely closed posteri- and sparser on pronotum), very finely and orly. Distance between mesocoxae about densely cellularly microreticulated to alu- 1.5 times and that between metacoxae taceous. Elytral surface with larger distinct about as great as that between procoxae. punctures (however smaller than eye fac- Metaventrite about as long as prostenum ets in diameter), interspaces between them and mesoventrite combined. Submeso- and with more relief microreticulation. Pygid- submetacoxal lines follow the posterior ium with sculpture and punctures as large edge of cavities. as those on pygidium, but intervals between Abdominal ventrite 1 nearly twice as them much smaller (about twice as great as long as hypopygidium and slightly shorter a puncture diameter). Prosternum with ob- than ventrites 2–4 combined. Epipleura at solete puncturation and nearly completely base slightly narrower than antennal club. smooth. Pterothoracic sterna and abdomi- Protibia somewhat wider than antennal nal ventrites with puncturation as that on club, subtriangular and with far projecting pygidium, but interspaces between punc- outer apical angle; meso- and metatibiae tures more or less smoothed. slightly wider and subtrapezoid, about 2.5 Head somewhat transverse, gently and times as long as wide, with slightly pro- evenly convex, with mediun-sized eyes jecting outer apical angle. Profemur nearly (composed by rather fine facets), lateral twice as long as wide, mesofemur about 1.5 lobes of frons forming arcuate convex curve; times as long as wide and metafemur about anterior median excision making visible 1 and 1/3 as long as wide; posterior edge of mandibular prosteca with setae. Labrum profemur rather deeply emarginate and that not exposed. Mandibles slightly exposed of meso- and metafemora nearly straight. from under frons and sharply curved along Tarsi very wide, stout and slightly lobed; outer edge. Antennae about as long as head setae of underside in tarsomeres 1–3 very wide; scape comparatively narrow, about reduced; ultimate tarsomere subflattened, three times as long as thick and slightly with excavated underside and simple nar- curved; 3-segmented club subcircular and row claws. composing a fourth of total antennal length, Aedeagus. Tegmen moderately sclero- antennomeres 9–11 comparable in length. tised, about twice as long as wide and Pronotum slightly transverse. Elytra wide- slightly emarginated at apex; penis trunk ly rounded at sides, their arcuate apices slightly sclerotised and with median exci- forming a very short and stump sutural an- sion at apex. gle, adsutural lines not expressed. Pygidium Etymology. The epithet of this new spe- slightly transverse. Anal sclerite does not cies means “most distinct”. exposed from under pygidial apex. DISCUSSION Mentum about 1.5 times as wide as long and with deeply and arcuately excised an- The new taxa here described demon- terior edge. Labial palpi with scarcely bend strate a rather great potential in variability palpomeres and with ultimate one about of some characters which are traditionally three times as long as thick. Ultimate max- used as diagnostic for known taxa of differ-

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(2): 274–298 296 A.G. KIREJTSHUK. “PARADOXAL” NEW GENUS AND SPECIES OF NITIDULIDAE ent taxonomic levels. Particularly, the diag- attempt is the generalisation was made by nostics of such large and diverse groups, as Audisio et al. (2009), in which the authors the genera Amphicrossus, Meligethinus and proposed many new taxa on the base of the Neopallodes here considered, gives an ex- cladogram first published in the mentioned pressive presentation of the problem. The paper. In contrast to the previous attempts, situation in larger taxa, as Epuraea Erich- this publication is not provided with either son, 1843; Carpophilus Stephens, 1829; Pria any explanation of the authors’ discrimina- Stephens, 1829; Meligethes Stephens, 1829; tion of the taxa, matrix or any list of charac- Mystrops Erichson, 1943; Camptodes Er- ters used for construction of this cladogram. ichson, 1943; Cryptarcha Shuckard, 1839; In general these authors seemed to use the Cybocephalus Erichson, 1844 and so on, grouping and characters traditionally re- many members of which still remain unde- garded as diagnostic for discrimination of scribed, is much more complicated. When the “species-groups”, which were treated as anybody includes in systematic study fossil preliminarily related by many researchers. sap beetles he encounter even with greater However, these groups of the genus Meli- difficulties in diagnostics and interpreta- gethes were rather frequently re-arranged, tion. Therefore the progress in understand- because there was not enough proper evi- ing of natural system and phylogeny of any dence if these preliminarily groupings fit group is somehow in significant dependence true relationship. on breadth of researcher views and standard The situation was somewhat improved of available knowledge. when the systematic studies on fauna of The new species of the genus Amphicros- the Palaeotropics became more intensive sus showed that this widely spread group and particularly when the comparison of instead of its probable antiquity still main- sequences of DNA of different species be- tains a comparative small level of differen- gan to fulfill. Nevertheless, these achieve- tiation and many peculiarities of its species ments represent only a beginning for such can be explained by homoplastic trans- large group as the subfamily Meligethinae, formations. Perhaps this situation could because many important species of it are be partly explained by some conservatism still undescribed and only for few species of the mode of life in species of this genus the molecular comparison was done. There- (most of them are associated with inhabit- fore the authors of the mentioned classifica- ance in tree sap and other liquid products of tion of the subfamily could not supply their plant origin under fermentation, although construction with a proper substantiation. some species are known as inquilines in The listed composition of many “genera” nests of ants). in it makes possible to suppose that the The new species of Meligethinus dem- authors meant to propose each “genus”, onstrates a mosaic of characters in the however to discuss this construction needs subfamily Meligethinae which are tradi- to obtain some more strict argumentation. tionally used as diagnostic for some gen- Some of the new “genera” proposed by the era. Endrödy-Younga (1978) undertook mentioned authors appear to be rather het- a first attempt of systematic revision and erogenous (e.g. Afrogethes Audisio et Cline “phylogenetic” reconstruction with cladis- in Audisio et al., 2009 or Indogethes Audisio tic methodology, results of which were es- et Cline in Audisio et al., 2009) and others sentially disproved by the subsequent ma- are certainly synonyms (e.g. Lucanopria terials on the subfamily. Later some such Audisio et Cline in Audisio et al., 2009 is a attempts were made and they were mostly very probable synonym of Microporum C. refuted after obtaining news data on the Waterhouse, 1876). Finally, it would be very groups considered or due to an increasing advisable to carry out more detailed studies a scope for consideration. The last large of the recent fauna of the Indo-Malayan and

Afrotropical Regions and particularly bring turelle in Paris and Smithsonian Institution). I the comparison of recent species into corre- have had also a support during some years from lation with the real fossil record. The con- the Programme of the Presidium of the Russian sidered classification of the subfamily Meli- Academy of Sciences “Origin of Biosphere and gethinae (Audisio et al., 2009) has not only Evolution of Geo-Biological Systems” and for ‒ scientific (methodological and conceptual) the work with the collection of ZIN from the Russian Ministry of Education and Science. Par- difficulties but also practical ones because ticularly, I greatly appreciate the help of A. Nel, in many cases users before identification T. Deuve, O. Montreuil, A. Mantilleri and other of generic attribution of any specimen are colleagues in MNHN, and also M.V.L. Barclay obliged to find its species attribution. Thus, and H. Mendel (BMNH), F. Hieke and M. Uhlig it is not reasonable to introduce this clas- (ZMB) for materials recent and fossil beetles, sification and the proposed names in taxo- and also for other various assistance during last nomical practice and it would be better for years. The preparation of drawings was made now to consider it as a preliminary proposal with assistance my daughter, Polina A. Kirej- till a publication of any substantiation of it tshuk. My sincere thanks also go to C.E. Carlton (Louisiana State University) for the possibility or at least a list of diagnostic characters of to study the holotype of Amborotubus clarkei Le- the “genera” to get possibily for evaluation schen et Carlton, 2004. I am also very grateful to of them for scientific use. S. Hisamatsu (Ehime University, Matsuyama) The new genus and new species of Cyllo- for his valuable remarks on an earlier version of dini and Cychramptodini show a very com- the manuscript. I also feel a pleasant duty to rec- plicated situation with these tribes. Some ognise a very creative editing and a patience to closer relationship of the Gonoglypha gen. the author’s opinions of the editor of this issue nov. from Australia to the rest members of N.G. Bogutskaya. the endemic Australian tribe seems to be more or less realistic. Nevertheless, some REFERENCES parallels in characters in this new genus and some groups of Cyllodini need to be stud- Audisio P. 1993. Coleoptera Nitidulidae – Ka- teretidae. Fauna d’Italia, 32. Bologna: Calde- ied in a more detail and explained in future. rini Edizione, i–xvi + 1–971 p. Probably the genus Gonoglypha gen. nov. Audisio P., Cline A.R., De Biase A., Antonini contains a mixture of apotypic and plesio- G., Mancini E., Trizzino M., Constantini typic characters, which will be important L., Strika S., Lamanna F. & Cerretti P. for a further phylogenetic analysis. Many 2009. Preliminary re-examination of genus- new genera and species of these tribes still level taxonomy of the pollen beetlesubfamily remain undescribed, particularly obtained Meligethinae (Coleoptera: Nitidulidae. Acta from the Neotropical and Indo-Malayan Entomologica Musei Nationalis Pragae, 49(2): Regions, including some forms from New 341–504. Guinea with pubescent dorsum. On the Cooper M.C. 1980. Species of the genus Meli- gethinus Grouvelle (Coleoptera: Nitiduli- other hand, the representatives of Cych- dae). Entomologica Scandinavica, 11: 32–36. ramptodini and Amborotubus could reach Endrödy-Younga S. 1978. Systematic revision some similarity in structure as a sequence of and phylogeny of some Meligethinae gen- similarity in their predaceous and inquilin- era from the Ethiopian Region (Coleoptera: ous mode of life. Nitidulidae). Entomologia Germanica, 4(3– 4): 295–316. ACKNOWLEDGEMENTS Jelínek J. 2000. New genus and species of Ori- ental Meligethinae with new observations Many scientific organizations supported the on the genera Cryptarchopria and Kabakovia study (a grant from the Russian Foundation of (Coleoptera: Nitidulidae). European Journal Basic Research 07-04-00540-а etc., some grants of Entomology, 97: 413–418. from the Royal Society, Deutsche Forschungs- Kirejtshuk A.G. 1994. Revision of the genus gemeinschaft, Muséum National d’Histoire Na- Neopallodes Reitter 1884 (Coleoptera Niti-

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