Nikolai B. Nikitsky
Generic classification of the beetle family Tetratomidae (Coleoptera, Tenebrionoidea) of the world, with description of new taxa
Pensott Series Faunistica No. 9
Generic classification of the beetle family Tetratomidae (Coleoptera, Tenebrionoidea} of the world, with description of new taxa
by
Nikolai B. Nikitsky
Sofia - Moscow 1998 ...PEJtSV E..I@Publishers
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© PENSOFfPublishers ISBN 954-642-039-5 Firstpublished 1998 All rights reseIVed
Editor: Dr. S. Golovatch Graphic and coverdesign: Dr.V. Milgram Onthe cover: Pisenuschujoi Miyatake, 1960 (original)
Printed by Biolnform SCIVices, Ltd., Moscow, Russia, January, 1998 CONTENTS
Ab stract ...... 5
Introduction...... 6 Mozphologicalc cs ...... 6 haracteristi gnosIS...... 10 Dia Key tosub families,tribes and genera of Tetratomidaebas edon imago ...... 10 Key tosub familiesand genera ofTetratomidae bas edon larvae ...... 14 SubfamilyTetratominae Bill berg,18 20 ...... 16 Genus TetratomaFabricius, 1790 ...... 16 Key tosubgenera of Tetratomabas edon imago...... 19
Tetratoma(Tetratoma ) tiiscoguttat.asp. nov ...... 21 Te tratoma(T.) nepalensissp.nov ...... 23 Te tratoma (T.) wittrJJ.erisp. nov ...... 25 Key to speciesof Tetratomabas edon larvae...... 26
SubfamilyPiseninae Miyatake, 19 60 ····························································�······· 27 Genus Pis enllSCasey, 1900 ...... 27 Key tospecies of Pis enllS basedon larvae...... 27 Genus No topisenllSNikitsky et Lawrence, 1992 ...... 29 GenusTri phylliaReitter, 1898 ...... 29 SubfamilyPentbinae Lacordaire, 1859 ...... 29
� Genus Cyanopenth c gen. nov ...... 29 Cyanopenthc sp. nov ...... 30 thailandica Genus Pcnth cNewman, 1838 ...... 32 Pcnth c reitterisp. nov ...... 32 SubfamilyHall omeninaeMulsant, 1856 ...... 33 Genus HallomcnllS Panzer, 1794 ...... 33 Ha llomcnllS (HallomcnllS} chinensissp . nov ...... ; ...... 35 HallomenllS (H.) orient.alissp.nov ...... 36 Key tospecies of Hall omenllS basedon larvae ...... 39 Genus MycetomaDejean, 1834 ...... 39
SubfamilyEustrop hinaeGistel, 1856 ························'.···················· ...... 39 TribeHolostrophini, tribus nov ...... 39 Genus PseudoholostrophllSNikitsky, 1983 ...... 40 Key tospecies of Pseudoholostroph11Sbased on imago ...... 40 PseudoholostrophllS (Holostrophinussub gen. nov. ) discolor (Hom, 1888), comb. nov ...... 43
Pseudoholostroph11S (Pseudoholostrophus) chinensissp. nov ...... 44 PseudoholostrophllS{P.)klapperichi(Pic, 1954) ...... 45
Pseudoholostrophus(P.) impressicollis(LeConte, 1874), comb. nov ...... 47 GenusHo lostroph usHom, 1888 ...... 48 4
Holostrophusminimus sp.nov ...... 48 Holostrophussimilis sp.nov ...... 51 Holostroph usorien. talis Lewis,1895 ...... 52 ?Holostrophusaureofasciatus(Pic, 1954) ...... 52
Key tospecies of Holostroph usb�ed on larvae ...... 53 TribeEustrophini Gistel, 1856 ...... 53 GenusEus trophusIlli ger,1802 ...... 54 Eustroph us)'UI1I111J1 en.sissp. nov...... 54 Genus SynstrophusSeidlitz,1898 ...... 58 Genus Eustrophopsis Champion, 1889 ...... 58 Eustrophopsis similis sp.nov...... 59 Eustrophopsis sexm.aculatasp. nov ...... 60 Eustrophopsis ochracea(Motschoulsky, 1872) ...... 63 Genus CurteustrvphinusPie, 1952 ...... 63 Acknowledgm.ents ...... 63
References...... 64
Index ...... 79 ABSTRACT
The beetlefamily Tetratomidae is diagnosed, withdetailed keys to subfamilies,genera and some species on TetratomaF. consideredas thesole constituent genus of based adults and laivae. is Tetratominaeincluding the subgenera Tetraromas. str.,AbstruliaCasey, Falsoxantbalia Pie, stat nov., Incolia Casey, nov. (bothpreviously regardedas genera), andParatetrafo 1111l stat full-rank subgen.nov. (type-species: Tetrafo1111lsabgutiiNakane,1955). new species described Three are genus: (T.) tiiscoguttBta nov. and (T.) nepalensis nov. fromNepal, and in this T. sp. T. sp. T.(T.) wittmeri sp. nov. from Bhutan. A new genus, Cyan.openth.e gen. nov., is established in the subfamilyPenthinae, with Penth.e me on, 1916 asits type-species and Cyan.openth.e taJlicaChampi nov., a,.q a second congener.Penth.e r:eitteri nov. descn"bed tha.il;mdica sp. fromThailand, sp. is from Sechuan, In thesubfamily Hallomeninae,which onlyincludes the genera HaJlomenus China. andMyceto1111l Dej.,the subgenus Parahaflomenus Nomuraet 1958, genusHall omenus, Pz. KatO, isconsidered as a new subjectivejunior synonym (syn. nov.) of thesubgenus XeuxesChampion, 1889, stat nov., thelatter tax.on previously regardedas agenus of full Basedrank. on a restudyof Pie, 1930, Hallomenus 1817, reticul1Jtus type specimens,XeuxeY diversicomis paJlensGyll.enbal, H Motschoulsky, 1872, H fuscosutumlis Blatchley, 1913 and H irmatusKangas, 1959 actually belongto neitherTetratomidae nor Melandryidae.Hall omenus variegatus Motschoulsky, 1872 and H 8D11Spioides Motschoulsky,1845 obviously do not belongin Tetratomidaeeither, but this could not reconfirmedby pertinent Twonew species described Hallomenus: be material. are in H chinensis nov. from Sechuan, China and orient.a1is nov. from India and sp. H sp. Kashmir, Pakistan.In the subfamily Eustrophinae,which includesthe genera Pseudoholostrophus Hazara, Nikitsky,1983, Holostrophus Horn,1888, Eustrophus Illiger, 1802, Eustrophopsis Champion, 1889 and S)'IlStrophus Seidlitz, 1898, a new tribe is established,Holostropbini tribus nov., to incorporateHolostrophusaruiPseJXkJholostrophU£ Holostrophusis typified forthe first time (type species: Eustrophus bifasciatus Say, 1824), with H minimus sp. nov. describedfrom Luzon, Philippinesand H similis sp.nov. from Sarawak,Malaysia. Based on a restudyof thetype of H multinotatus Pie, 1911, fromTaiwan, this taxon is considered as aninfraspecific color variation (syn. nov., stat nov.) of H orietJ.t.a1is Lewis, 1895. Eustrophinus aureofascliitus Pie, 1954, described from China,is transferredinto Holostrophus: ?H aureofasciatus(Pie, 1954), comb. nov. Pseudoholostrophusisconsidered to encompass two subgenera: Pseudoholostrophus s. str. with (Pie, 1954) (thetype-speci es, fromtype material ), impressicoJJis P. klapperichi redescribed P. (LeConte, 1874), comb. nov. ex Holo3trophus (redescribed) and P. chinensis sp. nov., from Sechuan, China, andHolostrophinus subgen. nov. for discolor (Hom, 1888), comb. erected P. nov. ex Holostrophus(redescn'bed.from a non-type).Eustrophus yunnariensissp.nov. isdescribed fromYunnan, Eustrophinus Seidlitz,1898 (type-specie5:Mycetophagus bicolor China. Fabricius, 1792), nov., is considered as a new junior synonym of Eustrophopsis Champion, 1889. syn. Eustrophopsiss . nov. andEustrophopsis se ulam nov. describedfrom imilissp xmac sp. are .la.ire and Mexico,respectively. Pseudore .hes'lli Fa.innaire,1883 (type-species:P. .niBJVS,ignam Fainnaire, 1883), thetype whichof could not beobtained for revision, is likely to represent a seniorsubjective synonym of Eustrophopsis,though no fonnalsynonymy isadvanced Basedon a restudy of here. the types of Eustrophus oc.bmceusMotschoulsky, 1872 and Eustrophinus bombinus Seidlitz, 1898,the speciesis considered as a new subjectivejunior synonym(syn. nov.) of theformer, latter Eustrophopsis ochracea (Motschoulsky, 1872), comb. nov. Curteustrophinus Pie, 1952 (type species: C overlaeti Pie, 1952), originallyin described Mel.anchyidae, isactually a member of Tenebrionidae. INTRODUCTION
For a longtime, the beetle family Tetratomidae Billberg, 1820 was oftenconsidered onlyas asubfiunily or tnoeo( or severaltribes within, the Melandryidae Leach, 1815 (e.g. Csiki, 1924; Kas7.ab, 1969; Arnett, 1973;etc.). Based on the Europeanfauna, Reitter(1911) referred toTetratomini, Eustrophini andHallomenini as independent tn'bes ofMelandryidae;we regardthese as subfamilies ofTetratomidae. Miyatake (1960) included the genera Tetrato maFabricius, 1790 andAbs truliaCasey , 1900 (both as members of thetribe Tetratomini), Pist:IJ. usCasey, I900 (as a member of thetribe Pisenini)and PCIJ. thcNewman, I 838(as a memberof thetribe Penthini) into the fiunily Tetratomidae.Crowson (1966) andViedma (I97I) consideredEustrophinae to be a betweenMelandryidae and Tetratom.idae. Hayashi (I975) referredHo lostrophus link Horn, 1888 toTetratomidae asbased onlarvalcharacters. wren La ce& Newton(l995) theTetratomidae subfamilies:Tetratominae, andPenthinae. divide into three Piseninae, We regardthe family Tetratomidae as particularly closely relatedto Mycetophagidae (Nikitsky, 1993), with the following genera involved: Tetrato maFabricius, 1790 (Tetratominae), Pist:nus Casey, I900, No topisCIJ. usNtlritsky& Lawrence, I992, and Triphyllia Reitter, I 898 (Piseninae) , PCIJ. thcNewman, I 838 and Cyanopt:D.thc gen. nov. (Penthinae), Hallomcnus Panzer, I 794 and Mycctoma Dejean, I 834 (Hallomeninae), Pscudoholostroph us Nik.its.Icy, I983, Ho lostroph us Hom, I888, Eustrophus Illiger,I802, Synstrophus Seidlitz,I898 andEustro phopsisC hampio� 1889 (Eustrophinae). To thebest of ourknowledge, one of the abovegenus-group names has neverbeen typified.Hence we designateits type-species forthe first time. Representatives of Tetratomidae occur almost all over the world except for the Australianregion with New Zealand.The world fauna amounts to about I 40 species.
Morphological characteristics
Imago. Length2-I 7 mm.Antennae I I-segmented, often with a sharplydefined 3- to 4-segmentedclub (Tetratominae,Piseninae) (Plate I: 8, I O; Plate 2: 6, 7, 9; Plate 3: I; Plate 4: 1, 9) or their3- 7 apicaljoints distinctly broadened (many of Eustrophinae) (Plate7: 10, 13; Plate8: 2, 8; Plate 9: 2, 4; Plate10: 7, 8; Plate 11: 8, 14), or antennae filiform(Plate 6: 7) or moniliform(Plate 6: I 9) (some Hallomeninae), or theirarticles serrate(Plate 6: I 5, I 7)(subgenus Xcux Cj,genus HaJJ omCD.us). Insome cases, antennae more or less filiform but withdistinct sex dimorphism(Plate 5: I, 3, 5) (PCD.thc), or witha serrateor pectinateclub (Plate5: 9-II) ( Cyanopt:nthc). Antennaeattached mostly freelyon each side of head, only in Piseninae their bases slightly coveredby lateral marginof frons.Mandioles apicallybidentate, either more or less symmetrical(e.g. Tctra toma) or noticeablyasymmetrical, in whichcase theirpreapical tooth or projection 7 of right mandible distinctly protruding(P late 10: 5) (e.g. Pist:D.us, HaJJomt:D.us, Holostrophus, Eustrophus). Chewingpart of mandiblesdistinctly protruding, mola can be well-developedand then with cleartubercles (Plate 10: 5) (e.g. Eustrophus). Galeausually considerably broaderand often even longerthan can be 1acinia,latter verynarrow (P late 10: 6)(e.g. Tetratomaand Eustroph us) . Lastsegment of maxillary palpsoften more or less oblong or of some other shape, elongate as a rule, suboval neithersharply norsecurifonn, usually either roundly or moreor lessstraight cultrifonn but obliquelytruncate so thatinternal side of joint shorterthan external one (Plate 1: 14; Plate 4: 6, 14; PlateS: 2; Plate 6: 2; Plate 7: 4; Plate10: 6). Prem.entum clearly separated from mentum, often sinuate (Plate 1: lS; Plate 10: 4) at anterior edge, sometimeswith a comb alongthe middle (Pist:D.us). Mentumclearly separatedfrom submentum,latter more ofteneven slightlyseparated from gula. Gula usuallymore if or less well-developed.Labial palps3-segmented (Plate 10: 4). Eyesmostly even if slightlysinuate (Plate S: 1, 3; Plate7: 1, 14; Plate8: 4), insome generavery large and almost contiguousupon frons (Plate 11: 5) (Synstroph us, manyof Eustrophopsis). Frontoclypeal suture often more or less well-noticeable (Plate 11: 5). Head behind eyesusually without nec k-like,sharp constriction. · Pronotumclearly transv erse,considerably wider than head (P late 1: 1; Plate3: 1; Plate 4: 1, 12; PlateS: 1, 9; Plate6: 1, 8; Plate7: 3; Plate8: 3; Plate 10: 3), withlateral edge bordered, oftenanterior edgeand atleast partly basebordered as well. Pronotal sides not :frequentlyrounded, itsdisk with neitherribs nor elongatestriae (excluding basal impressions). Pair of basalimpress ions on pronotumoften more or less well developed(Tetratominae, Penthinae, Hallomeninae, Pist:D.us and partly Eustrophinae) (Plate1: 1; Plate 3: 1; Plate 5: 1, 3, 9, 10; Plate 6: 8, 18; Plate 7: 3; Plate 8: 1). Scutellum.always Elytrapresent fully developed, more or less parall el-sided, weakly broadenedposte1forly or oblong-oval,distinctly narro wedposteriorly (Plate 7: 12; Plate 8: 1, 3, 7; Plate 9: 3; Plate10: 3). Elytraconfused ly punctured(T etratominae, Piseninae, Holostrophini, partlyHall am.co.us andEustro phopsis, tosome extent Cyanopenth.c) or rowsof punctures,the number of rows exceoiing11 in (Plate5: 1). or withstriae PCD.tht: Procoxae considerably 1ransverse, as a rule, somewhat projecting, div ided by prostemalprocess, latterwell-visible in situ(without special preparati on), even apices ofprocoxae not contiguous(Plate 2: 8, 13; Plate7: lS; Plate8: 9; Plate9: 5; Plate 10: 14; Plate 11: 7, 10, 13). Trochantin ofprocoxae canbe well-dev eloped(Plate 2: 8, 13) (at leastTetratominae, Piseninae, My ct:fomaand partlyHaJJ omco.us). Anteriorcoxal cavitiesopen from behind. M esocoxaediv ided,mesepimera reachingthe mesocoxal cav ities(Plate 10: 13; Plate11: 3). Metastemumusually well-developed, not shorter thanvisible abdominal stemite 1. Metepistemawhole or wi1han anteriorly more or less well-separated,not rarelysomewhat elevated, triangular area developed best in Penthinae andEustrophinae (Plate 10: 13; Plate 11: 3), sometimes to some extent noticeablealso in Hallomeninae.Metacoxae strongly 1ransverse, simple (always without narrow, triangularpiece separatedby suturefrom posterior part of coxa,punctured in a differentway, directed obliqu ely anterolaterally). Legs moderatelylong; tibiae usually 8 not or onlyslightly shortened; metatibiae at least slightly longer than tarsomere 1 (Plate 1: 1; Plate4: 1; Plate 5: 1; Plate6: 8; Plate7: 3; Plate10: 15). Meso-andmetatibiae can beartransverse rows of spines at outer edge (Eustrophus, Eustrophopsis). Spurs of metatibiaeoften fairly short, only ratherseldom ea 1/2 lengthof tarsomere1. Tarsi of alllegs simple (Plate 1: 1; Plate 4: 9; Plate 5: 1; Plate6: 1, 18; Plate 10: 15), without sharplybroadened, lobuliformjoints, pattern 5-5-4 in bothsexes. Claws simple andat most only somewhat thickened at base. Wmgs more or less well-developed, more oftenwith well-developed radial cell, M-Cu-loop, 1-2 analcells and not rarelywith a more or less well-developed subcubital fleck (Plate 1: 2). In Hallomeninae and Eustrophinae, 1st analcell canbe opener closedby only a weaklydeveloped piece of vei n (Plate 9: 11; Plate 11: 2). Metendostemite more or lesswell-developed, with a marked stem,often with (Tetratominae, Penthinaeand Ho lostroph us) (Plate Piseninae, 4: 15; Plate9: 12) or withoutlateral plates directed outside(Plate 6: 3; Plate 10: 18). Abdomenwith five visible stcmites. Abd ominaltergite 8 moreor lessrounded at apex (Plate 1: 18; Plate 10: l); true stemite 8 often sinuate(Plate 10: 1, 11), in Tetratoma witha ratherthin, medial, sclerotized process in basalpart (Plate 1: 17). Genitalsegment oftenwith more or less well-developed,scleroti.zed lobesin anteriorpart, these lobes usuallynei separated, thervery strongly narro w nor long,more or lessrounded at anterior edge(Plate 10: 2, 12); morerarely, genital segment more or lessunilobate (Plate 9: 16) or withvery long andnarrow lobes (e.g. Synstrophusmacro phthalm us(R.eitter, 1887)) (Plate10: 16). Scleroti.zedrods in posteriorpart of genitalsegment coming together or close toeach other caudally, moderately to fairly long (Plate 10: 2, 12), theirposterior extent canparallel or virtuallycontiguous. Aedeagus of varyingstructure , oftensimple, witha pairof elongatedp arameresand a more or less elongatedpenis (T etratominae, Penthinae, Triphyllia, Synstrophus,Eustroph us,Eustroph opsis)(Plate 1: 5, 6, 9, 11; Plate4: 17; Plate 5: 4; Plate 11: 11, 12), orparamerestransformed into lobes of more complex structure, long, not :frequently setigerous, strongly divided (Myccto� Hallomt:IJ.us,Ho lostroph us) (Plate6: 10, 20; Plate8: 6). InHolostrophini,penis usually of complicated �tructure, underside with a scleroti.zed process connected by thin processeswith penial apex, latter bearing thin spinif ormprocesses (Plate 7: 8; Plate8: 6; Plate 9: 9); parameresofHolostrophini highly moveablyconnected with a strongly elongated basal part of tegmen. Sometimes (Pist:IJ.us) basal part of tegmenbearing eithera pairoflong sensorysetigerous lobes on ventralside (Plate4: 2, 4, 7) or only one shorterlobe (NotopisCD.us) (Plate4: 11 ). Ovipositor membranous in apical part, oftenwith more heavily sclerotizedlateral areasand a strongly elastic medial part (Plate 9: 17). Stylestactile, supplied with elongate setae at apex, as arule.
Larva. Length4-18 mm. Body often more or lessconvex and parallel-sided (Plate 14: 1; Plate17: 1; Plate19: 1; Plate21: 1). Headinm ost generawith a moreor lesswell developedepicranial stem, with five ocelli on eachside ofhead,as arule (Plate 12: 3; Plate 14: 4; Plate 15: 1; Plate 17: 5; Plate 19: 7). Hypostomalrods usually shortor of 9 medium (Plate14: 3; Plate19: 3). distinctlyseparated clypeus, from more or siz.e Labrum lessstrongly rounded at frontedge (Plate 14: 6; Plate15: 3) or som sinuate(Plate ewhat 12: 5; Plate21: 6). Sclerotizedrods of epipharynxeither nearly undeveloped or short (Tetratominae,Hall omeninae,Eumophinae) (Plate 12: 5; Plate14: 6; Plate17: 4; Plate 18: 4) torelatively long (Penthe, Hayashi,1972) (Plate 16: . often after 7).Antermomere l rathershort and 1ransverse, more rarely about ss longas broad; 2nd one longerfor the most moreor less longitudinal or about as long as 3rd6ne considerably part, broad; nmrowcr1han2nd(Plate14: 5;Plate15: 4;Plate16: 5;Plate17: 7;Plate21: 10); seosorimn of anteonomere2 moreor less longitudinaland conical, asa role,varying in lengthin relationto aotennomere3. Mandibles (exceptT etiatominae), preapical tooth asymmetrical orprojection on rightmandible usually developed stronger 1han the on left one (Plate14: 7;Plate15: 2;Plate17: 2;Plate19: 4). Molarpartwell-developedinPiseninae(Plate14: 7; Plate 15: 2) and,to some extent, inMyceto ma(Plate 17: 2); in Tetratom.inae and Eustrophinae,part chewing of mandibleswith a prosthecoidstructure or prostheca,not rarelylike thin, short, transparent processes (Plate 12: 2; Plate 19: 4). InPentinae and Ha/Jommus,prosthecaandmolaeithernotmarlced orlatterpoorly-developed.Maxillae witha well-developed areaand a theirmalamorestrongly orless articular cardo, rounded atanterior edge, obtuseor moreor lessstraight not falcate(Plate 12: 1, 6; 11'1mcatebut Plate14: 3, 8; Plate 15: 5; Plate16: 3; Plate 17:3; Plate19: 3; Plate21:9).Anterior and inner edges of mala can bear thickened setae or spines (Piseninae, Penthinae, Hallomeninae)(Plate 14: 8; Plate15: 5; Plate17: 3), or inneredge without such setae/ spines (Tetratominae, Eustrophinae)(Plate 12: 8; Plate19: 5; Plate21: 9). Maxillary palps3-segmented, their joint 1 ratheroften more or lesstransverse and shorter than 2nd, latteroften more or less longitudinalor about aslong asbroad (seldom transverse),not rarely considerably longerthan lst;joint 3 longitudinaland often considerably longer that2nd. Labimnconsisting of twoparts separated by sutures: a transverseprementum andamore strongly elongated m entumclearly separatedeither from subm.entmn or from subm.entum+ gula(Plate14: 3; Plate15: 5; Plate16: 3; Plate19: 3). Ligulausually more or lesswell-developed, rather short and rounded at apex (considerably shorter than labial palps)(Plate 12: 1; Plate14: 3; Plate17: 3;Plate19: 3; Plate21:7); the only exceptionis Pmtb.e whichhas a long(not shorterthan palps labial ), moreor lessparall el-sidedligula (Plate16: 3). Hypopharyngealscl eromacan be more or lesswell-developed (especially so in Piseninae,see Plate 14: 9; Plate15: 5). Legsfrom rclati velyshort medium.-six.ed; to coxaeusually narro wly separatedand more or less strongly projecting, between distance themconsiderably shorter than coxal transverse diameter, asa role;claws in most cases clearly curved andusually with two setae of unequallength (Plate 12: 7; Plate14: 11; Plate 15: 6; Plate 17: 8; Plate 9: 10). Posterior parts of minaltergites 2-8 atleast abdo eitherwith one well-developed,row transverse of elongatedsetae (Plate 13: 5; Plate14: 1; Plate17: l;Plate18: 6;Plate19: 1; Plate21:l) orwith microscopicalspinules(Pmtb.e) (Plate16: 1). Anterioredge of scutumof atleast:preapical tergitesnot frequently abdominal with a transverse sclerotizedline. Abdominal tergites usually withoutwell-developed 10
tergite8 notlonger1han 7tb, tergite 9 of usual, structure ambulacralampullae. Abdominal witha pairofurogomphi at apex withoutsclerotized fovea them(Plate 13: but between 1, 5; Plate14: 13; Plate15: 7; Plate16: 15; Plate17: 9; Plate 20: 1, 3; Plate21: 1). Abdominal tergite 9 oftencovered with numeroustubercles sclerotized (Piseninae, I:IallOID.eninae,Eustrophinae)(Platel 4:13,16;Plate15:7;Platel 7:9;Platel 8:9;Plate
20: 1,3; Plate21: 1 ). Inneredge of urogomphi not rarely with well-developed setigerous tubercles (Tetratominae) (Plate 13: 1, 9), sclerotized teeth or lobes (Piseninae, Holostrophini)(Plate 14: 13;Plate 15:7; Plate 20:3, 5).hl.E ustroph opsis,urogomphi can sclerotizedtubercles. Abdominal segment 10 usuallyposteroventral or ventral bear (Plate 14: 14; Plate20: 2). Spiraclesoften ann ular-biforous(Plate 14: 15; Plate16: 17; Platel 7:ll ;Plate 19:ll ;Plate21:13). Developmentmostly invarious xylotrophic fungi, main ly Aphyllophorales.
Diagnosis
TheTetrato midaediffers from the Melandryidae, to which for a long timeit belonged as a subfiunily(see above), as follows.The imago ofTetratomidae characterizedby is thesimple narrow of tarsi alllegs, as :well as by theprocoxae separated by a prostemal processcombined with the simple metacoxae of usual structurethe and relatively short spursofthe metatioiae(not longerthanhalfoftarsomerel but mostly not morethanl/3, as a rule).The prementmn, mentmn and submentmnof thelarvae are oftendivided by suilJres abdominal tergite 9 alwayswith a pairof urogomphi but without sclerotized , fovea in between. Such a combination of characters usually absent from the true is Melandryidae(Nikitsky, 1989a).
Key to subfamilies, tribes and genera of Tetratomidae based on imago
1. Each elytrumwith over 11 rows of punctures(Plate 5: 1 ), if punctationmore or lessconfused, then antennae with a pectinate4- or 5-segmentedclub (Plate 5: 9,
11) . Body broad (Plate5: 1, 9, 10), not or weaklynarro wedposteriorly, not less than 9in mm length.Upperside black, metallic dmk blue or violet,sometimes with a more brightly colored scutellum. Scutellum almost always(except P.pimelia (Fabricius,1787)) withdense light reddish or yellowishpubescence contrasting againstbackground of elytra(Plate 5: 1, 3). Antennae more or less filifo� but theirstructure and proportions at leastof some antennomeres( 4th-7th)in males andfemales considerably different (Plate 5: 1, 3, 5),or antennaewith a pectinate club. Pronotmnvery strongly transverse(Plate 5: 1, 3, 9, 10),usually not lessthan 2 timesas broadas long, not rarelywith light pubescence in posteriorpart before 1 1 scutellumand with two very strongly developed basalimpressions. Trochantin of procoxae well-developed. Metepistemawith a well-separated, not frequently somewhatelevated, triangular areain anterior part (Plate7: 5). Metendosternite withlateral plates (Plate5: 6).Wmgs with radial celland two closedanal cells. Aedeagususually of simple structure (Plate 5: 7, 8), witha pairof distinctly divided param.eresand a simple penis...... (subfam.Pen.thinae) ... 2 -. Eachelytrum with not morethan 11 rowsof punctures (Plate 6: 18; Plate10: 15), ifpunctation confused, then. antennae always without pectinate club. Body oblong oval, eitherdistinctly narro wed posteriorlyor lessthan 9 mm. Scutellumusually withoutdense light ccntrasting pubescence. Male and female antennae similar in structure, either almost the same or very slightlydifferent in proportions of antennomeres (Plate 6: 9, 19; Plate 8: 2, 8; Plate 10: 7). Pronotumoften more weaklytransverse, than less 2 timesas broad as long, itspair ofbasal impressions can be weaklydeveloped ...... 3 2. Antennaefiliform in bothsexes, but withdistinct sex dimorphism.in structureof medialant ennomeresat least(Plate 5: 1, 3, 5).Body black, usually with a clarified
lastant ennomere(Plate 5: 5) ...... Penthe'Newm. -. Antennae in females at least (as we know allspecies of this genus solely 5-om females)with a serrateorpectinate 5-segmented 4-or club (Plate 5: 9, 10). Upperside metallic dark blue or violet, often with more brightlycolored scutellum. Last
antennomerenot clarified...... Cyanopenthe gm nov. 3. Antennaewith a 3- or 4-segmen.ted abruptlyclub defined (Plate 2: 6, 20; Plate4: 1, 13). Elytiaconfusedly punctured, body gen erallymore stronglyparall el.:.sided. ofprocoxaeusuallywell-visible. Metepistemawithout triangular Trochantin isolated areain anteriorpart. Metendostemite with lateral plates directed0utside (Plate 4: 15), wings often. withmore or less well-developedradial cell and two analcells. Spursof metatibiae short (Plate 4: 9) ...... 4 -. Antennaefiliform (Plate 6: 7), serrate (Plate 6: 17), moniliform(Plate 6: 19), or apicaljoints moreor lessgradually or rather weaklybroadened (Plate7: 7, 10, 13; Plate10: 7, 8) (in lattercase, with a well-definedtriangular area in anteriorpart of metepistema). Body often more stronglyoblong-oval and stronger narrowed posteriorly...... 7 4. Antenna!club 3-segmented (Plate 4: 1, 9, 12). Sutureseparating frons from clypeus semi-circularor bow-shapedand more weakly impressed. In tionaddi to parameres, aedeagus not rarelywith setigerous sensorylobes (Plate 4: 2, 4, 11) ...... (subfam.Piseninae) ... 5 -. Antenna!club 4-segmeoted, though antennomere 7 canbe fairly strongly broadened too (Plate2: 6, 7, 9, 12; Plate3: 1). Suture separatingfrons from clypeus close to straight and stronglydeveloped. Aedeagus simple, with a pair of separated parnmeresand a more or less stronglyelongate penis (Plate1: 5, 6, 9, 19) ...... � ...... (subfam.Tetratominae) . .. Tetratoma F. 5. Antenna! club large and strongly 3symmetrical, not shorter than preceding antennomerescombined (Plate 4: I 0). Aedeaguswith one unpairedsensory lobe 1 2
(Plate4: 11) . Pairof basal foveaeon pronotumnot developed(Plate 4: 9). Lateral edgeof pronotumsomewhat reflexed and flattened. lst visible abdominalstemite considerablylonger than 2nd but much shorterthan 2nd and 3rd ones combined.
Uppersidecovered with somewhat raisedhairs (Plate 4: 9) ...... No topisetJ. usNikitsk:y et I..a wrence -. Antennalclub considerablyshorter than preceding ant ennomeres combined,more or less symmetrical (Plate 4: 13). Aedeagus with (Plate 4: 4) or without pair of
sensorylobes (Plate4: 17) ...... 6 6. Pairof basalimpressions onpronotum eitherundeveloped or verysmall, puncture like and not frequently hardly visible (Plate 4: 12). Lateral edges of pronotum distinctlyflatten ed. st1 visibleabdominal stemitenot or veryslightly longerthan
2nd one. Aedeaguswithout sensoryprocesses (Plate 4: 17) ...... Triphyllia Rtt.
-. Pairof basalimpressions on pronotum not largebut well-developed (Plate 4: 1 ). Lateraledges of pronotumnot flattened(Plate 4: 1 ). lst visible abdominalstemite usuallymuch longer than2nd. Aedeagus with a pairof sensorylobes (Plate 4: 2,
4, 1) ...... PisetJ. us Casey 7. Antennae filiform, serrate or more or less moniliform (Plate 6: 7, 14, 15, 19). Pronotum with 2-3 well-developed fovea-like impressions (Plate 6: 8, 12, 18). Metepistemawhole or weaklydivided into two pieces: a short triangularanterior pieceand an elo ngate-quadrangularposterior one. Metendostemitewithout lateral plates directedoutside. P arameres of aedeagus lobe-shaped, strongly separated
(Plate 6: 5, 10, 20) ...... (subfam..Hallomeninae) ... 8 -. Apical 3-7 antennomeres usually more or less considerably broadened, at least some of these transverse (Plate 7: 7, 10; Plate 8: 2, 8; Plate 9: 4; Plate 10: 8). Pronotumwith two veryweak (Plate 10: 3, 15) or stroke-likeimpre ssions(Plate 7: 3, 9) in basalpart Metepistemadistinctly subdivided into a shorttriangular anterior and anelo ngate-quadranguposterior piece (Plate 7: 5). Metendosternitewitho ut lateralplates (Plate 6: 3) outside directed(excluding Holostrophini).Wmgs more oftenwith a clearly closed anal cell and usuallyonly with trace of a crosspiece closing anotherce�1 ...... ( subfam.Eustr ophinae) ... 9 8. Pronotum with 3-4 strong impressions (Plate 6: 18); elytrausually with 10 (sometimes partly confused) rows of punctures;space between lst and2nd rows of punctures broadened at least in basal part of elytra and coarsely punctate.
Antennaemonilif orm(Plate 6: 19). Trochantinof procoxaewell-visible ...... My ccto111Jl. Dej . -. Prunotumonly withtw o strongbasal impressions (Plate 6: 8, 12). Elyt:raconfusedly
punctured or with less than 10 weaker rows of punctures (Plate 6: 1 ). Antennae filiform (Plate 6:7), more or lessmoniliform or serrate(Plate 6: 15, 17). Trochantin of procoxaenot or more weaklydeveloped ...... HallometJ.usPz. 9. Meso- andmetatibiae without transverserows of spines atouter edge(Plate 7: 3). Elytra confusedlypunctured (Plate7: 12). Eyes considerablyseparated (Plate 7: 1, 1 3 14; Plate8: 4). Metendostemitewith lateral plates (Plate 9: 12).Te grnenand penis of complexstructure , latterwith a spine-likestructure anterior at edge (Plate 8: 6). Pronotum.more oftenwith a pairof more or lessstroke-like impress ionsin frontof base(Plate 7: 3, 9; Plate 8: 1, 7). Elytradark in the most part, witha redspotty pattern,more rarelyone-color (Plate 7: 3; Plate 8: 3, 7) ...... (tribeHolostrophini, trib. nov.) ... 10 -. Meso- andmetatibiae with transverserows of spines at outer edge(Plate 10: 1 O; Plate11: 3 ), ifwithout them (Synstro phus) , theelytra with rows of punctures(Plate 10: 15) andeyes almost contiguous. Metendosternite usually without lateral plates directedoutside (Plate 10: 18). Aedeagususually with apic allydivided p arameres of simple structureand with penis (Plate 11: 11, 12) lackinga complexspine-like
structure...... (tri"beEustrophini) . .. 11 10. Eyesmore stronglysinuate and more stronglyapproximate each to other, distance between eyesless than ocular transverse diameter (Plate 8: 4). ProsterruJprocess usually strongly surpassing posteriorof edge procoxae(Plate 8: 5; Plate 9: 5).
Elytra(excluding H. unicolor Lewis,18 95) darkwith a redpattern ...... Holostrop li.usHo m -. Eyes more weakly sinuate and more broadly separated, distancebetween eyes somewhatexceeding transverse ocular diameter (Plate 7: 1, 14). Prostemalprocess mostlymargined on eachside andnot or onlyrather slightly surpassing posterior edgeof procoxae(Plate 7: 2, 11). Elytraone-color, usually red-brown, brownish or blackwith a lighthumeral spot (Plate8: 1) or witha vaguetransverse fascia in basal (Plate 7: 12) ...... Ps eudoholostrophusNikitsky part 11. Meso- andmetatibia e without transverserows of spinesat outer edge (Plate distinct 10: 15). Eyesvery large, strongly appro ximateto each other, alm(Jst contiguous on frons;distance between eyes not less than7-8 timesshorter than ocular transverse diameter.Elytra with rows of puncturesor striae. Apical 6- 7 antennomeresdistinctly broadened. Pronotum strongly transverse, with a small, usually with apair of shallow basalimpressions (Plate 10: 15). Transversefold/sut ureof hypomeronof prothoraxin its posteriorpart from almost tomore or lesswell-developed indistinct ...... Synstrophus Seidl. -. Meso- andmetatibiae with well-developed transverse rows of spines atouter edge (Plate llJ: 10; Plate 11: 3) ...... 12 12. Hypomeronwithout distinct transverse fold/suture in itsposterior part (Plate 10: 14). Eyes more or lessconsiderably separated, distance betw eeneyes usually not more than2 timesless than ocular transverse diameter (Plate 10: 3). Elytrawith rowsof puncturesor striae(Plate 10: 3). Baseof pronotumoften margined at least partly.Apical5-7 antennomeresdistinctly broadened(Plate 10: 7, 8). Pronotum
stronglytransverse, with a pairof weak impressions (Plate10 : 3) ...... b$3.l ...... Eustrophus ill. -. Hypomeronwith a distincttransverse fold/suture in itsposterior part (Plate 11: 3). Eyes verystrongly approximate toeach other, as arule; distance between eyes 14 considerably over 2 times less than ocular transverse diameter (Plate 11: 5). Antennae with 5-7 enlarged apical joints (Plate 11: 4, 8, 14). Posterior edge of prosternalprocess strongly sinuate to al.most straight cate(Plate 11: 7, 10, 13). trun Elytrawith rows of punctures,as arole (Plate 11: 6, 9), but sometimesmore or less
confusedly punctured(Plate 11 : 1) ...... Eustrophopsis Champ.( = Eustrophinus Seidl., syn. nov.)
Key to subfamilies and genera of Tetratomidae based on larvae
1. Epicranial stem not developed (Plate 21: 2) or extremely short. Mandibles asymmetrical,with prostheca consisting of rathernumerous seta-likeprocesses thin (Plate 21: 5) (accordingto Viedma, 1971, can be a modifiedmolar thiscture stru partof themandi"bles ). Labrumfairly deeplyemarginate (Plate 21: 6).Abdominal tergite9 with numeroussclerotized tubercles, latter can be present also inbasal ofurogomphi (Plate 21: 1, 12). Endocarina well-developed (Plate 21: 2). half Spiracles biforous(Plate 21: 13) ......
...... (subfam.Eustrophinae, in part)(Plate 21: 1-13) ... Eustrophopsis Champ. -. Epicranialstem well-developed (Plate 12: 3; Plate 14: 2; Plate 15: 1; Plate 16: 2; Plate 17: 5; Plate 18: 1; Plate 19: 2). Innerpart of mandiblesof different structure (Plate12: 2; Plate 14: 7; Plate 15: 2; Plate16: 8; Plate17: 2; Plate 18: 2; Plate19: 4). atmost shallowly sinuateat anterior edge,usually straightor rounded Labrum (Plate 12: 5; Plate 14: 6; Plate 15: 3; Plate16: 7; Plate17: 4; Plate 18,4; Plate 19: 6). Basal ofurogomphiusually without or witha few sclerotized tuberclesat half leastdo rsally(Plate 13: 2, 3, 6; Plate 14:17; Plate15: 8; Plate 16: 15; Plate17: 10;
Plate18: 7, 10; Plate20: 6) ...... 2 2. Abdominaltergite 9 withoutnumerous sclerotized tubercles on dorsal surface (their numberusually not exceeding6-8) (Plate13: 1, 3, 5; Plate 16:15). Epicranial stem usually short(over 5 timesas short ashead length up toanterior edge of labrum) (Plate 12: 3; Plate 16: 2). Mandibleswithout well-developed mola (Plate 12: 2;
Plate 16: 8) ...... 3 -. Abdominaltergite 9 withnumerous sclerotized tubercles on dorsalsurface (Plate 14: 13, 16; Plate15: 7). Epicranialstem better developed, not more thantimes 5 as shortas head length (Plate 14: 2; Plate 15: 1). Mandibles not rarelywith a mola (Plate 14: 7; Plate 15: 2) ...... 4 3. Ligula roundedapically, apparently shorter thanlabial palps (Plate 12: 1, 8). Mala withneither spines nor setaeat inner edge, withsolitary setae or bristlesat macro most (Plate 12: 1, 8). Mandibles moreor less symmetrical,with a prostheca-like structure(Plate 12: 2).Abdominal tergitestransverse without rows of microspinules, at most with a erse sclerotized line in their anterior part (Plate 13: 5). transv Urogomphi withwell-developed setigerous tubercles at inner edge (Plate 13: 1, 4,
5, 9) ...... (subfam.Te tratominae).. . Tetratoma F. 15 -. Llgulalong, more or lessparall el-sided,umally not shorterthan labial palps(Plate 16: 11). Maia withwell-developed thickened setae at inner edge(Plate 16: 9). Mandiblesfairly sharply asymmetrical (without prostheca-like structure), left one witha well-developedsharpened pro jectionnear middle (Plate16: 8). Abdominal tergiteswith microscopic denticles (Plate 16: 1, 14). Urogomphi withoutwell-
developedsetigerous tubercles at inner edge(Plate 16: 15) ......
...... (subfam.Penthinae) ... Pen.th.eNewm. 4. Urogomphi witha distinctdenticle or processon theinner side (Plate14: 13, 16; Plate15: 7; Plate 20: 1, 3, 5). Epicranialstem usually somewhat shorter, not more than1/3 lengthof cephalic capsule(measuring its lengthup to base oflabrum) (Plate 14: 2; Plate19: 2). Mandibles with a more or less well-developed mola (Plate 14: 7; Plate15: 2) orprostheca-likestrocture(Plate19: 4) ...... 5 -. Urogomphi with neitherdistinct denticle nor process on inner side (Plate17: 9; Plate18: 6, 9). Epicranialstem somewhat better developed, usually over 1/3 length of cephaliccapsule (Plate 17: S; Plate 18: 1) ...... (subfam.Hall omeninae)... 7 5. Mandibles without distinct mola, anterior part of their chewing surface with a prostheca-like structure(Plate 19: 4). Hypopharyngeal scleroma not Qr weakly developed.Sensorium of antennomere2 not morethan half long as asant ennomere 3(Plate 19: 8) ...... (subfam.Eustrophinae, in part) ... HolostrophusHom -. Mandibleswith a mola (Plate14: 7),but canbe without clear transverse distinct they rows ofmicrotubercles on ventralside (Plate 15: 2). Hypopbaryngealscleroma well-developed(Plate 14: 9; Plate15: 5).Sensorium of antennomere2 overhalf as long as 3rd(Plate 14: 8; Plate 15: 4) ...... (subfam.Piseninae) ... 6 6. Baseof innerpart of urogomphiwith a rathershort, sharp denticle, latter not less than3-4 times as short �s urogomphi inlateral view (Plate 14: 14). Urogomphi properfairly thin lateral in view, not less than2.0-2.5 timesas long asthick at base
(Plate14: 14, 17) ...... Piscn.usCasey -. Base of inner partofurogomphi with a fairly long lobe, latter 1.3-1.S times as shortas urogomphiin lateral view (Plate15: 8). Urogomphiproper thick in lateral view,over most of extentof anterior edge fairly straight,unciform at apex, 1.3-1.4 timesas long asthick at base (Plate 1 S: 8) ...... Trip hyllia Rtt. 7. Epipharynx over most of its extent along middle with well-developed rows of microtrichiae(Plate 17: 4). Mandibleswith a more or less well-developed mola butwithout transversemicroscopic rows of tubercles on ventralside (Plate 17: 2).
Labrumless strongly transverse (Plate 17: 4) ...... MycetomaDej. -. Epipharynxwithout well-developed rows of microtrichiae(Plate 18: 4). Mandibles without mola (Plate18: 2). more stronglytransverse (Plate 18: 4).. . distinct Labrum ...... Hallom en.usPz. 16 Subfamily Tetratominae Billberg, 1820
Generaincluded: TetratoI111lF. only.
Genus Fabricius, 1790 Tetratoma
Plate 1: 1-23; Plate 2: 1-24; Plate3: 1-5 Type-species: TetratoI1111.fimgorum. Fabricius, 1790 (Plate 1: 1, 2) Modemdescrip tionsof genuscan be foundelsewhere (Nikitsky, 1989a, 1992), this in theabove family characteristi cs and keys� well asin thedefinitions of thesubgenera givenbelow. Theworld fauna encompasses 22 species,all nicely correspondingto the scopeof theHolarctic. Tetrato I1111.hasbeen encountered in Nepaland Bhutan, Himalaya forthe time,though it long known Sikkim ( cyanoptera Champion, first has been from T. 1924). Larvaas in Plate12: 1-9; Plate 13: 1-9. Some authorsdivide taxoninto two genera(e.g. Csiki,1924; Arnett,1973) or this subgenera (Crowson, 1964; Nikitsk:y, 1992): TetratoI1111. F. and Abstmlia Casey. Division of genusinto subgenera s eemsto be useful,but we also include therein this the earlier monotypic genus Fcls oX1lllthalia Pie, 1934, stat nov. (type-species: F. bicoloripes Pie, 1934, describedfrom in thefamily ofTenebrionidae ), theearlier China monotypic genus Incolia Casey, 1900, stat nov. (type-species: Incolia Jongipemiis Casey, 1900, from NorthAmerica), and ParatetratoI1111. subgen. nov. (type-species: Te tratomasakagu tiiNakane, 1955, fromJapan). Thesubgenus Ab stmlia Casey,1900 di:fers from Tetratoma s. str.in combining thefollowing characters: antenn omeres8-11 shorteror than about long as asall precerling ones combined(Plate 2: 20, 21 ); parameres verynarro wly divided or contiguousall along inneredge, sharpened at apex. Penismuch shorterthan tegm.en (Plate 1: 11, 12, 19, 23) (this feature makes all the species of Abstrulia we know from males distinguishablefrom other subgenera); lateraledges of pronotummore oftenmore or less unevento undulate-crenulate,usually fairly broadly reflexedand flattened (Plate 1: 1O; Plate 2: 22); 1 stvisible abdominalsternite oftenconsiderably longerthat 2nd (Plate 1: 13, 16, 20), 5thone withan impression at least in males.Elytra bicolor, black or brown, with a red spotty pattern (Plate 1: 10; Plate 2: 23). Upperside distinctly thoughnot rarelyrather shortlypubescent. subgenus overthe Holarctic and includes the foUowing species: This is distributed T.(A.) anco .raFabricius,1790 (Plate1: 19; Plate 2: 22, 23), T.(A) virgoMotschoulsk:y, 1845(b), (A .) co Baudi, 1877, (A .) bau eri P , 1864, (A.) ainu T. creni llis T. du erris T. (Nakane, 1963) (Plate 1: 23; Plate 2: 20), T. (A .) japonicaMiyatake, 1955 (Plate 2: 21), T. (A .) pictipemiisReitter, 1896, T. (A .) tcdaldiReitter, 1887, T. (A .) tesselata Melsheimer,1844 (Plate 1: 10-12), T.(A) variegata(Casey, 1900). Thespecies of this subgenuswe met in nature more oftendevelop on Stereaceaeor Corticiaceaefungi. 1 7
1 21 8
.. Q14 --·17 - � AC �"Q> 15 11 • 2 ��· 10 '\) · . 5· 9 ;;] �3 _19 . . ' � . t=�16� . 03 7 22 4
P�te 1. General view and details of structure of Tetrat.oma spp. (5, 6, 11, 12 - afterMiyatake, 1960, other.1 original). 1, 2 - Tetratoma (Tetrat.orna)fungorom; 3-T.(T.) cyanoptera (type); 4 - T. (Incoli;J)con color; 5 - T.(T.) nobu chii; 6 - T.ma) (Paraletrato 7 - T. (Falso.ranthalia)bicoloripcs T.(F.) desm1m:stii; 10-13 - T.(Ab struliJJ) sabgutii;14-19 - T.(A.)an co..ra;20, 23 - T.(A.) (type);8-9 - tesselam; ainu; 21-22 - Tetrat.oma(Incoli;J)Jongi pemiis. 1, 8, 10, 21 - generalview; 2 -wing; 3, 7 - pronotum;
4, 13, 16, 20, 22 -visible sternites 1 and2 of abdomen; 5, 6, 9, 19, 23 - aedeagus(dorsal view); 11 - tegmen; 12 - penis; 14 - joints 2-4 of maxillarypalps; 15 - labium; 17 - abdominalsternite 8; 18 - abdominal tergi.te 8. 1 8 The subgenus TctraloI118s. str. characterizedby theant enna!club morestrongly is elongate in comparison withthe flagellum together with joint 1 (antennomeres 8-11 longerthan all other joints combined) (Plate 2: 6; Plate3: 1) . Parameresusually more broadly separated,if narro wly separated,then not sharpenedat apex; penis fairly long, always in relationto tegmenlonger than in Abstrulia(Plate 1: 5; Plate 2: 1, 4). Lateral edges of pronotumoften more even. 1st visibleabdominal stemite about aslong as 2nd (Plate1: 4), 5thone withoutimpression. Upperside with very short pubescence, lookinglike almostglabrous. Coloration of elytraoften of metallictinges (dark blue or green),more rarelywith reddisha spotty pattern. subgenus representedin the Holarctic by thefollowing species: TetratoI118 This is (TetratoI118) fimgorumFabricius, 1790 (Plate 1: 1, 2), T.(T. ) tnmcorumLeConte, 1879, T.(T. ) talyshens.tSNikitsky, 1989, T.(T. )nobuchiiNakan.e, 1955 (Plate 1: 5), T.(T. ) cyanoptera Champion, 1924 (Plate 1: 3; Plate 2: 4-6), T. (T.) fuscoguttatasp. nov. (Plate2: 1-3; Plate 3: 1, 2), T.(T. )nepalens.tSsp.nov. (Plate 3: 3), T.(T. ) wittmerisp. nov. (Plate3: 4, -5). Based on the main features given above for the other subgenera, the subgenus Fa lsoX11IJ.thaliaPie, 1934, statnov., closerto Tdralc1I18s. str. Itsantennomeres 8-11 is arelonger than all preceding ones combined (Plate 1: 8; Plate2: 9). However,the main differenceof Falso xantha/ia.from. Tctratoma s. str.lies in thewell-developed, moderately densepubescence on thedorsal side of thebody, whereeach preceding seta distinctly surpassesthe base of thenext one (Plate1: 8). Only two Palearcticspecies: T.(Falso X11IJ.thalia) de.miarestiiLatreille, 1807 (Plate 1: 8, 9), which distributed overEur ope and theCaucasus, and (Falsoxanthalia) is T. bicoloripes(Pic, 1934), comb. nov. (Plate 1: 7; Plate 2: 9-11), described andknown only fromGinfushan, Theformer species hithertobeen treated as a member China. has of Te trato1111l s. str. The subgenus IncoliaCasey, 1900, statnov. (type-species: 1 JongipennisCasey, 1900)has heretofore been considered as a genus of seemsto takea position full rank. It intermediate between the above subgenera as based on the following characters: antennomeres8-11 slightly shorterthan or aslong as allpreceding joints combined (thisfeature brings it nearerto Abstrulia(Plate 2: 16, 17)). Pubescenceof elytrarather short, decumbent,developed weaker than in Fa lsoXJlilthalia(Plate 1: 21), but stronger than inTetrato1111l s. str. Sides of pronotumconsiderably reflexedand can be uneven ( T.(/.). Jon.gi pt:DIJis(Casey), see Plate 1:21) oralmost even with lateral edge somewhat protruding laterally nearmiddle (T. (lncolia) concolor LeConte, 1879). lst visible abdominalstemite almost not or slightlylonger than (Plate 2nd 1: 22), 5thone without fovea.Aedeagus apic allywith distinctly divided p arameresand with a long penis; this makes subgenusmore similarto Tetrato1I18s. str.(Plate 2: 18, 19). Uppersidecan this be more or less darkone-<:olor or bicolor. Two species can be included in subgenus: (lncolia) longipennis(Casey), this T. comb. nov. (Plate1: 21-22; Plate 2: 17, 19) and T.(lnc olia) concolorLeC.(Plate 1: 4; 19 Plate 2: 14, 16), bothfrom NorthAmerica. The latterspecies has hitherto beentreated as amemb erof Tetratoma s. str. The Japanesespecies Tetratomasabgutii Nakane, 1955 (Plate 1: 6; Plate2: 12), knownto us from a single male only, is here considered as representing a separate subgenus of its own, Paratctratomasubgen. nov. , whichdiff ers thefrom othersubgenera in the following combination of characters: strongly developed antenna! club eres8-1 1 considerablylonger than onescombined); very narrow (antennom allpreceding (8-10 times asnarro w as transversediameter of coxa) prostema!pr ocess separating procoxae (Plate 2: 13), a feature absentfrom other Tetratoma species, in which it is considerablybroader; almostglabrous elytra; metallicblue dark or greenuppers ide; andwell-developed medialimp ressionon visible abdominal5 stemite ofmale (Plate 2: 24), the latterfea ture sharedonly with the species of the subgenus Abstrulia.
Key to subgenera of based on imago Tetratoma
1. Penisconsiderably shorter than tegmen, para.meres narrow at apex, more or less pointed and usually more or less contiguous (Plate 1: 11, 12, 19, 23). Visible abdominal stemite5 witha ratherwel l-developedimpression. Ant ennomeres 8-11 considerablyshorter than or aboutas long asall preceding ones combined (Plate 2: 20, 21). Upperside moreor lessdistinctly (setae pubescent well- visible in dorsal view), elytrablack or black-brownwith a red-yellow, redor yellow spottypattern (Piate 1: 1 O; Plate2: 23). I.ateralsides of pronotumoften uneven fairlyand strongly
re:tlexed...... Subgenus Abstrulia Casey -. Penisnot orvery slightly shorter thantegmen (Plate 1: 5, 6; Plate2: 1, 2, 4, 10, 14, 18, 19). Para.meres apically usuallydistinctly separated, if almost contiguous,then not sharpened (Plate 1: 5, 6; Plate 2: 1, 10, 14, 18). VIS ible abdominal stemite 5 (excluding T.sakagu. tiiNakane) withoutwell-developed fovea. Ant�omeres 8- 11 usually longer(rarely slightly shorter or equal - subgenus Incolia, see Plate 2: 16, 17) thanall precedingones combined (Plate 2: 6, 7, 9; Plate 3: 1) ...... 2 2. Antennomeres8- 11 considerablylonger than all precedingones combined(Plate 1: 1; Plate 2: 6, 9; Plate 3: 1). Upperside almost glabrous (setae very short and hardlyvisible) or pubescence fairlystrongly developedand eachpreceding seta
distinctlysurpassing the base of nextone ...... 3 -. Antennomeres 8-11 slightlyshorter than orlong as asall preceding ones combined (Plate 2: 16, 17). Pubescenceof upperside distinct, but rather short, withwhich eachpreceding hairusually not reachesbehind a base of thenext (Plate 1: 21). Lateralsides ofpronot ummore orless clearly re:tlexed(Plate 1: 21). Elytrawithout markedred-yel low spots againsta dmk background, atmost with a clarifiedbasal part. 1 st visible abdominal atsternite most only slightly longerthan 2nd (Plate 1: 22). Species fromNorth America ...... SubgenusInco lia Casey 20
2 3
12 20 7 21
23
· . ...
. -�. .�' _,:·•··.-...,_
I . \ ; 4 \. ( 16 18 19 )0 XY 13 8 24
Plate 2. Generalview and details ofstruc tureof Tetratoma spp. (23 - afterKasz.ab, 1969, 4, 7, 12, 14, 15- afterMiyatake, 1�60, othersoriginal) . 1-3 - Tetratoma (Te tratoma) fuscoguttata (type); 4-6 - T.(T.} cyaaop tem(type);1- T.(T.} nobuchii;8- T.(T.} fimgomm; 9-11- T.(Falso xanthalia) bicoloripes (type); 12, 13, 24 - T. (Pa ratctratoma) salaJgutii; 14-16- T. (lncolia) concolor; 17-19 - T. (lncolia) longipeDIJia; 20 - T. (Abstrulia) ainu; 21 - T. (A.)jap onica;22, 23 - T. (A.) ancora. 1, 18 - tegm.en (dorsal view); 2, 15, 19 - penis (dorsal view); 3 - penis (lateral view); 4, 10, 14 - aedeagus (dorsal view); 5, 11 - apex of penis (lateral view); 6, 7, 9, 12, 16, 17, 20, 21 - antenna; 8, 13 - prostemal process; 22 - side edge of pronotum; 23 - various types of elytral pattern; 24 - visible abdominal stemite 5 of male. 21 3. Prostemalpr ocess separatingprocoxae verynarr ow, 8-10 timesas narrow ascoxal transverse diameterof coxa (Plate 2: 13). 5thvisible abdominalstemite at leastin male witha strongimpre ssion (Plate 2: 24). Sides ofpronotum rather smooth and weakly reflexed. Upperside almost glabrous, metallic blue, dark blue-green or darkblue . Known onlyfrom Japan ...... Subgenus Para tetratoma subgen. nov. -. Prostemal process broader, usually not more than 5 times as narrow as coxal transverse diameter (Plate 2: 8). 5th visible abdominal stemite without fovea. Upperside either almostglabrous but sharplybicol or, or fairly stronglypu bescent andmore uniformone-color ...... 4 4. Upperside rather densely andmore or less long pubescent, comparatively one color (Plate 1: 8); such combinations of characters as red pronotum andmeta llic darkblue, greenor sharplybicolor elytraabsent ...... SubgenusFal soxanthalia Pie -. Uppersidealmost glabrous, setaevery shortand can be hardly seenin dorsal view. Elytrasharply bicolor (black or brownwith a red spottypatt ern) (Plate 3: 1, 3, 5) or metallicdark blue, green, darkblue-green, violet or black,often in combination withar ed pronotum...... Subgenus Te tratoma F.
Te tra toma {Tetratoma) fuscogutta ta sp. nov.
Plate 2: 1-3; Plate 3: 1, 2 · Holotypemale, Nepal, ParbatDistr., Ridge E. GhoropaniPass, 3100 m, 7 .X.1983, Smetana& LOblleg. - 5 paratypes, including 1 female, same locality; 3 paratypes: Nepal, ParbatDistr., Ghoropani Pass, N. slope, 2750 m, 5.X1983, Smetanaet Llibl leg. Theholotype and most of theparatypes are kept in the collection of the Museum d'Histoire naturelle in Geneva, while 2 paratypes in the Zoological Museum of the State Universityof Moscow, Russia. Description. Head black withyellow-brown clypeus andlabrum. Antennomer es 1-4 or 1-7 reddish-brown, other apical antenna! joints infuscate; pronotum usually red-brown,only its sides canbe somewhat clarified.Stem.um and abdomenusually dark brown-blackwith somewhat clarifiedsides of prosternum and abdomen. Scutellum dark. Legs red-brownto brown. Elytrayellow-brown, shining, at base with a black brown or black longitudinal spot located medially of humeral angle, and with a transverserow ofthree spots in anterior 1/3 extent (inner spot reaching the sutureand outer spot connectedwith in:fuscatean lateral margin); apical partof elytrablack-brown, entirely margining a large transverse yellow-brown spot. Elytra with very shortand Sparsegrayi sh hairs not rarelyhardly distinguishab le in dorsal view. 22
Antennaerather short, not swpassing 116 extent of elytra, theirjoi nts 8-1 1 1.2- 1.3 times longer thanall preceding antennomeres combined (Plate 3: 1 ); antennomere 1 more or less roun� longitudinal; 2nd somewhat narrower and ea. 1.4 times shorter than lst, also somewhat rounded; antennomere 3 elongate-triangular, 1.1-1.2 times longerthan 2n� 1.6-1.8 times aslong as broad and asantennomere 4, latter about as long asbroad or very slightlytransverse; antennomere 5 ea. 1.1-1.2 times broaderand longer than4th; antennomere 6 a little shorterthan 5th and 1.25-1.3 times asbroad as long; antennomere 7 ea. 1.7- 1.8 timesas broad aslong and 1.3 times broader than6th; antennomere 8 ea. 1.2-1.3 times broader and 2.2-2.4 times longer than 7th, 1.1-1.2 times as long as broad; antenna! joints 9 and 10 about as long and broad as 8th; antennomere 11 more or less oblong-oval, 1.5-1.7 times longer than both wide and antennomere 10. Head shining, somewhat flattened, with dense, relatively fine punctation, distance between punctures considerably less thantheir diameter. Vertex straight behind eyes not frequently with a more or less strong impression. Width of :frons between distinctly sinuate eyes 1.2-1.3 times greater than transverse ocular diameter. Pronotum shining, not shagreen� 1.5-1.6 times asbroad asboth long and head with eyes. Sides ofpronotum more orless straight, reflexedand flattened not toobroadly, 2-4-undulate, hind comers of pronotum rectangular, not frequently somewhatrounded orslightly sharplyprotrudi ng. Greatest width of pronotum in its anteriorpart, whence naITowedmore or less roundly anteriorly and oftenvecy weakly rectilinear posterio rly. Base of pronotum in front of scutellum weakly sinuate. Disk of pronotum with two postmedialimpressions on each side behind middle and withtwo well-developedbasal fqveae. Punctation of pronotum dense, of rather medium size, distance between punctures considerably less thantheir diameter. ,� Scutellum ea. 1.4-1.5 times as broad as long, shining, finely and rather sparsely punctured. Elytra more or less oblong-oval, shining, virtually not shagreened, considerablybroadened posteriorly, ea. 1.6 times aslong asgreatest width, withdense andrough punctation (considerablycoarser than on pronotum), in middle partof elytra distancebetween neighboring punctures 1.5-3 .0 times less thandiameter of a puncture. Prosternal process posteriorly vecy weakly broadene� ea. 3 times as narrow as coxal transverse diameter. Mesostemalprocess considerably broader thanprosternal one and reaching the middle of length of mesocoxae. Metastemum shining, not shagreen�with dense and rather coarsepunctati on.Abdomenwith finer, rather sparse minal than punctation. 1 st visible abdo stemite almost not longer 2nd; stemite 5 without impression. Aedeagus as in Plate 2: 1-3. Body length: 3.8-5.0 mm. Diagnosis: The spottypattern of the elytra coupled with theirvecy short dust-like pubescence, as well as the strongly developed antennomeres 8-11 which are longer than all preceding joints combined make this species safely distinguishable from congeners. 23
Te tra toma (T.) nepalensis sp. nov.
Plate 3: 3 Holotype female,Nepal, Ramche, 1800-3350 m, 18. VI.1978,Bhakta B. leg. Thetype is kept in theNatu.rh istorisches Museumin Basel. Description. Underside, also including legs, clypeus, labrum, three basal antennomeres, pronotum(except fortransverse row of obscure darkspots) andelytra (except for dark spots) yellow to yellow-brown or red-yellow-brown; upperside iridescent; scutellum light, more or less dirty yellow-brown; antennomeres 4-7 considerablydarker than preceding ones, antennaljo ints 8-11 black-brown; head dark re d-brown; disk of pronotumred-y ellow-brown with atransverse row of fourblack brown obscure spots, of which inner longitudinal and outer one transverse; black brown patternof elytra consisting of a spot bilobed in posteriorpart at base, one spot common to bothelytra on suture behind scutellum, a transverserow of three spots in frontof middleof elytra,with outer spot connectedwith lateral edge of elytrabroadly infuscateto up apex; apicalpart of elytrabla ck-brown, margining two relativelysm all light spots, of which one, more distantfrom apex, largerand more stronglyapp roximate tosut ure thansides, to while theother small er, apical,lon gitudinal, considerablymore strongly approximate to suture(Plate 3: 3). Body pubescence asin preceding species. Antennomeres 8-11 less than 1.2 times as long as allpr eceding ones combined. Antennaljo int 1 broad,roundly triangular, almo st aslong asbro ad; 2nd considerably narrower and only slightly shorter thanlst; 3rd narrow, elongate-triangular, ea. 1.3 timesas long as2nd and1.8-1.9 times aslong asbr oad; antennomere4 about aslong as broad andea. 1.6 timesas short as 3rd; antennaljo int 5 somewhat rounded,about as long as 4th and not more than 1.05 times as broad as long; antennomere 6 slightly shorterthan 5thand ea. 1.3 timesas broad as long; antennomere7 aboutas long and ea. 1.2 times as broad as6th, 1.5 times as broad aslong; antennaljoint 8 ea. 1.3 times as broad andea. 2.0-2.2 timesas long as7th, about as long asbro ad andbarely longer thanantenn aljoint 9 or 10; antennomere 11 oblong-oval,ea. 1.5 times aslong aswi de andas lOth. Headshining, denselybut not coarselypuncture d; width of fronsbetween eyes ea. 1.25 times greater thantransverse diameter of a distinctly sinuateeye. Pronotumshining, stronglytransverse, ea. 1.6 times asbro ad aslong and1. 7 times as broad as head with eyes. Base of pronotum weakly rounded, only very weakly . sinuate in frontof scutellum ; sides of pronotumnarr owly reflexedand flattened, not entirely even, each with two very weak, roundly angulate projections, in between almost parallel-sided, posteriorly narrowed more or less rectilinearly and anteriorly narrowed roundly; hind comers obtuseand rounded; anteriorpart of medialsurf ace of pronotum with a longitudinal impression flanked by a rounded impression situated more closely to posterior part of pronotum; deeply impressed areas of paired basal impressions of pronotum transverse, while theirshallower areas roundly triangular and narrowed anteriorly. Disk of pronotumwith dense, moderately coarsepunctation, 24
5
3
�0
4 1 2
Plate 3. Generalview and detailsof structureofT etratoma spp.(all original). 1, 2-Te tratoma (T.) fuscogutf1lta; 3 - 5 - 1 - general view; 2, 4 -punctation of elytra T.(T.} nepalensis; 4, T.(T.) wittrri.eri. (x 20); 3 - pronotum and elytra; 5 - elytrum. 25 distance between punctures on the average considerably less than their diameter. Posteriorpart medialof surfa ce of pronotumwith a shortened smoothpatch. Scutellumweakly transverse, shining, with rathersparse punctation of about same size as on pronotum. Elytra more or less elongate, shining, considerably broadened posteriorly, ea. 1.2 times as broad as pronotum and 1.55 times as long as their own greatest width; theirpunctation very rough, considerably coarser thanon pronotum, distancebetween punctures in middle partof elytra not more than 1.5 times less than diameter of a puncture. Boey length: 3.8 mm. Diagnosis: The species differs from T..fiis coguttata, sp. nov. in coloration of the upperside(Plate 3: 3), lighter legs and less strongly elongated antennomeres 8-11.
Te tra toma (T.) wittmeri sp. nov.
Plate 3: 4, 5 Holotype male, Bhutan, Chimakothi, 1900-2300 m, 14.5.1972, Bhutan Zool. Exp. leg. The type is kept in thecollection of theN aturhistor isches Museum in Basel. Description. Most ofhead andmaxillary palps darkbrown-red, antennomeres 8-11 black-brown, basal ones distinctly clarified. Underside red-brown; clypeus, labrum, legs (with somewhatinfuscate tarsi) and elytra (excludingdark spots) yellow toyellow brown; disk of pronotum infuscate, red-brown, with darker two, thinsemi-arches in medial partof pronotum, with a short, small lineiformpatch behind thesesemi-arches in themiddle, and aninfuscate area ou tside. Scutellumbl ack-brown. Each elytrumwith a black-brown longitudinal spotat base; anobliquely transverse spot near suture in anterior 113; a small longitudinal spotsituated somewhat outside and behind preceding spot; a horseshoe-shaped spot laterad of preceding spot andreaching the infuscate lateral edge (latterbroadly continuedup to apex of elytra); a largeirr egular diamond-shaped spotstarting before and ending up far behind middle of elytra, yet at a considerable distance from their apex; and also a dark longitudinal stripe starting from the darklateral spot situated straight behind themiddle of elytra andcontinued backwards almost in parallels toinfusca te lateraledge of elytra but reaching neithertheir apexnor suture, nor diamond-shaped spot (Plate 3: 5). Body pubescence as inpreceding species. Antennaenot surpassingbasal 1/4 extent of elytra, antennomeres 8-11 ea. 1.1-1.15 times as long as allpreceding joints combined; antennomere 1 large, distinctly longitudinal, 1.4-1.5 times longer and considerably broader than a rounded 2nd, latter 1.2 times aslong asbroad; antenna!jo int 3 elongate-triangular, 1.3-1.4 times as long as 26
2nd and 1.8 times as long as broad; antennomere 4 somewhat rounded, ea. 1.7-1.8 times as short as 3rd and about as long asbroad; antennaljo int 5 ea. 1.5 times as long and 1.1-1.2 times asbroad as4th, about aslong as broad; antennomere 6 considerably shorter than 5th and ea. 1.3 times as broad aslong; antennaljoint 7 almost not longer but ea. 1.3 times broader than 6th, 1.6 times asbroad as long; antennomere 8 ea. 1.25 times broader than7th, 1.2 times as long as both preceding joints combined, ea. 1.1 times as long as9th or 1 Oth,both latter subequal; antenna!joint 9 weakly longitudinal, 1.1-1.2 times aslong as broad; antennomere 11 oblong-oval, ea. 1.5 times longer than 1 Oth and 1.7 times as long as broad. Head shining, not shagreened, with dense, not coarsepuncta tion; widthof frons between distinctlysinuate eyes ea. 1.5 times greater thantransverse oculardiameter. Pronotum transverse, 1.6 times broader than head with eyes and 1.55 times as broad aslong; its lateral sides slightly uneven, very weaklyrounded and rather weakly reflexed, anterior comers of pronotum entirely rounded, posterior ones more or less rectangularly rounded; pronotum broadest near middle whence more strongly and roundly narrowed anteriorly thanbasally. Medialpart of pronotum beforemiddle with a longitudinal impression. Punctation of pronotum rough and very dense, punctures than separated narrby very ow, small,wrinkle-like spaces,latter by farnarr ower diameter of a puncture. Scutellumweakly transverse,rather sparsely and moderately finelypunctured. Elytra oblong-oval, ea. 1.55 times as long as broad and ea. 1.2 times broaderthan pronotum, distinctly broadened posteriorly, their surface convex, shining, with very dense and roughpunctation, considerably coarserthan on pronotum, distance betweenpunctures in middle partof elytral disk3 .0-3 .5 times less than diameter of a puncture. Aedeagus similarto thatof T.fusc oguttata sp . nov.
Body length: 3.6 mm. Diagnosis:This species differs fromT. fiis coguttata sp. nov. and T.nep alensis sp. nov. in combining such characters as the veryrough anddense punctation of theelytra (Plate 3: 4), black-brown scutellum andcoloration of the upperside (Plate 3: 5).
Key to species of Te tra toma based on larvae
Plate 12: 1-9; Plate 13: 1-9 1. Urogomphi more narrow and proportionately elongate, notch between them at level of inner setigerous tubercles considerably greater than width of one urogomphus together with tubercles (Plate 13: 5, 9). Medialtubercles of abdominal tergite 9 more narrow and weakly developed (Plate 13: 5). Body only with slightly
yellowish scuta oftergites. Body length: 4.3-5.5 mm...... (Subgenus AbstruliaCasey) ... 2 -. Urogomphi broader andproportionately short, notch between them more shallow, distance at level of inner setigerous tubercles not or very slightly exceeding the 27 widthof one urogomphustogether with tubercles (Plate 13: 1, 4). Medial tubercles of abdominaltergite 9 broaderand more stronglydeveloped (Plate 13: 1, 4). Scuta
of abdominal tergites notrarely considerably darker. Body length: 6. 5 - 8.5 mm3 2. Inner setigerous tubercles of urogomphi more strongly shiftedventrad, usually only slightly projecting inside in dorsal view and thus more broadly separated
(Plate 13: 5) ...... T. (A bstrulia) ainu(Nakan.e) -. Innersetigerous tubercles of urogomphimore stronglyproj ecting and thusmore
stronglyapproximate to eachother (Plate 13: 9) ...... T. ?4-) an.con F. 3. Abdominaltergite 9 in lateral view bearinga transverse row of 2-3 large setigerous tubercles with smaller ones in front of them (Plate 13: 3). Medial tubercles of
abdominal tergite 9 considerablylarger, directed laterad (Plate 13: 4) ...... T.(Falsoxa n.thalia) desmarestii Latr. -. Abdominal tergite 9 in lateral view bearsnot more than2-3 tubercles (Plate 13: 2). Medialtuber cles of abdominal tergite 9 considerably more weaklydeve lopedand
directed caudad (Plate 13: 1) ...... T. (Te tra toma)fimgomm F.
Subfamily Piseninae Miyatake, 1960
Genera included: PisenusCas ey, 1900, No topisenusNikitsky et Lawrence, 1992, and Triphyllia Reitter, 1898.
Genus Pisenus Casey, 1900
Plate 4: 1-8 Type-species: Cryp tophagus hum.eralis Kirby, 1837 Thisgenus is represented in the world fauna by six species, of which three are distributed in the south of the Russian FarEast, in Korea and J.apan, one species in Taiwan andtwo in NorthAmeri ca. Larvaas inPlate 14: 1-18.
Key to species of Pisenus based on larvae
1. Teeth at base of interior edge of urogomphi more elongate and sharp, 1.7-2.0
times aslong as broad at base (Plate 14: 13, 14) . Body larger: 6.0-7 .5 mm......
...... 2 -. Tubercles at base of interior edge of urogomphi less elongate andmore strongly rounded, not more than 1.4 times as long as broad in basal part (Plate 14: 16).
Body smaller: 4.0-5.0 mm. Kunashir Island;Japan ...... , ...... P.insi gnis (Reitter, 1889) 28
17
2 5
3 '
� : . .. : •· ' :· , . ···· ·'
... . ,. . ·· .I : . I 7
11
15
14 12
Plate General view and details of structure of Piseninae(1-5, 7, 8 - after Miyatak:e, 1960, others 4. original). 1-3 - Pisenuscb ujoi; - P.rufitarsis; 1, 8 - insignia; 9-11 - No topisen usboleti; 12-17 - 4-6 P. Trip .hyUia .koenigi l, 9, 12 - general view; 2, 7 - tegm.en (ventral view); 3, 5, 8 - penis; - 4, 6, 14 maxillary palp; 10, 13 - antenna; 11, 17 - aedeagus; 15 - metendosternite; - wing. 16 29
2. Antennomere 3 usuallyshorter than 2nd (Plate 14: 5). Khabarovskand Maritime provinces, Sakhalinand Kun ashirislands; Korea, Japan...... P.mfitarsis (R.eitter, 1 889) Antennomere 3 not shorter, rather often even somewhat longer, than2nd (Plate 14: 18). Khabarovskand Maritime pro vinces, Sakhalinand K1mashir islands; Korea, ...... chujoi Japan P. Miyatak:e, 1960
enus No topisenus Nikitsky et Lawrence, 1992
Plate 4: 9-11 Typ e-species: No topisenusboleti Nikitsk:y et Lawrence, 1992 (Plate 4: 9-11) This monotypic genus has been described from Chile and still is the only representative of the subfamily Piseninae occurring beyond the true Holarctic (considering Taiwan as a territory intermediate between the Holarctic and Oriental regions).
Genus Triphyllia Reitter, 1898
Plate 4: 12-17 Type-species: TriphyUia koenigi Reitter, 1898 (Plate 4: 12-17) This genus includes two species, one of which is distributed in the Northwest Caucasus, aswell as inWest andcentral Transcauc asia, andthe otherin NorthAmerica.. The genusEupis en usCase y, 1900, whichuntil recently encompassedthe Nearctic E. elongatus (LeConte, 187 5) (see Nikitsk:y, 1988), can now be considered only as a subgenus of the genus TriphyUia. Itdisplays fuller wing venation (with a developed radialcell) and the aedeagus carries a group of setae missing in the Caucasian Triphyllia koenigiReitter, 1898. Larvaas in Plate 15: 1-9.
Subfamily Penthinae Lacordaire, 1859
Genera included: Cyanopenthe gen. nov. and Penthe Newman, 1838
Genus Cyanopenthe gen. nov.
Plate 5: 9-11 Typ e-species: PenthemetaUica Champion, 1916 (Plate 5: 10, 11) 30 Cyan openthediff ers from Penthein the following characters: apical 4-5 antennomeres, all stronglybroadened asymmetrically, more or less pectinate-clavate; lastantennomere not lighter thanpreceding ones (Plate 5: 9, 11 ), andbody coloration of metallictin ge. Only femalesare curr entlyknown. Two species. Distribution:India, Bhutan and Indochina.
Cy anopenthe th ailandica sp. nov.
Plate 5: 9 Holotypefemale, NW Thailand, Doi Suthep, 19-23.04.1991, Jan Farkac leg. The typeis kept in thecollection of theNaturhi storisches Museum in Basel. Description. Upperside shining, not shagreened. Antennae, femora, tibiae and undersidevery darkblue, partly sternum and abdomen with violet tinge; head and pronotum more or less black, fronsat antenna!bases and pronotal edges withme tallic darkblue lustre, elytraviolet with darlcbluish suture (partly) and lateralScutellum edges. metallic green, with dense decumbent yellow pubescence almost undevelopedin its medialpart against backgroundof a darkerimpre ssion ofblack-violet color. Upperside withblack elongate protrudingpu bescence. Joint 2 ofmaxillary palps elongate-tri angular, subequalto 4th,latter almost twice aslong asbroad, sidesof 4th more or less parallel, apexobliquely roun ded.Antennomere 1 broad, strongly longitudinal, ea. 1.4 timesas long as2nd, lattershort and very weakly longitudinal; antennomere 3 strongly elongate, ea. 4 times as long as broad and ea. 3.4-3.5 times longer than 2nd; antennal ea.joint 4 1.6 times asshort as 3rd,2.5 timesas longas broad andsli ghtly longer than latter5th, about twice as long asbr oad; antennal joint 6thlongitudinal, ea. 1.4 timesas long asbroad, slightly shorterand cons iderably broaderthan 5th; antennomeres 7-11 broadened into a pectinateclub; antennal joint7 aboutas long as 6th, 1.8 times as broad as long and 2.2-2.3 times broader than 6th; antennaljoints 8-10 each aboutas long and 1.4 times as broad as 7th,2.4-2.5 times as broad as long; antennaljoint 11 asymmetrically curved, oblong-oval, ea. 1.8 timesas long asbroad. Head shining,more or less convex,with fineirr egularpunctation.Frontal widthbetween eyes ea. 1.3 times less thantransverse ocular diameter. Pronotum shining, strongly flattened (ea. 1.8 times broader than head with eyes and2.4 timesas broad as long), withfine sparse punctation (distance between punctures near middle ofdisk equalto 2-3 diameters of a puncture). Edging of pronotum well developed, reflexedon eachside andin lateralparts of anterioredge; posterior edge on eachside (approximatelyto level of inner edgeofbasal impressions) broadly margined. Anterior ofedge pronotum near middle very shallowly sinuate. Pair ofbasalimpressions ofpronotum very large and extendingfrom baseto about 112length . Pronotumbroadest near posterior comers, its sides more or less parallel posteriorly, but more or less rectilinearlyand somewhat roundlynarr owedanteriorly. 31
10
3
.·
5
t� ��,, t; ;:• !�, .. ...
.
� 9 I� � . I' 6 7 8
Plate 5. General view and details of structure of Penthinae original). 1, 2 - Pcnthejapana (all Marseul, 1876 (male); - P. reitteri (male); 5 - P. mfop ubcns Marseul, 1876 (female); 6 - P. 3, 4 obliquata; 7, 8 - P. almoreIJSis Champion, 1917; 9 - Cyao.opcntbe thailandica 10, 11 - (type); C. metJJUica (type). 1, 9, 10 - general view; 2 - maxillary palp; 3 - anterior partof body; - aedeagus 4 (dorsal view); 5, 11 - antenna; 6-metendosternite; 7 - tegmen (dorsal view); 8 - . penis 32 Scutellum large,transvers e, densely punctured, ea. 1.6 timesas short as pronotum. Elytrabroadly oval, shining, generally more or less convex, somewhat flattened on disk, at humeri ea. 1.3 times asbroad aspronotum at base, posteriorly considerably broadened, ea. 1.55-1.6 times as long as broad. Rows of punctures of elytra rather roughbut clearly traceable onlyin middle partof disk of each elytrum, while towards edges andsuture theirconfiguration becoming more or less confused. Widthof intervals betweenrows ofpunctures upto 0.3-0. 7 diameterof punctures in rows. Spaces between rough rows of punctures with dense, rather fine punctures, latter' s diameter 2.0-2.5 times smaller thanthat of puncturesin rows but considerably largerthan punctures in thecenter ofpronotal disk. Prostemum before procoxaeea. 1.5 times as short ascoxal longitudinal diameter. Prostemalpr ocess moderately stronglybroadened posteriorly (on each side without distinct edging frombehind) , nearmidway ea. 2.3 times as narrow as coxal transverse diameter. Posterior, more or less roundly truncated part of prostemal process not surpassingposterior edge of coxae caudally.Metepistema shining, withfine punctation of medium density. 1 st visible abdominal sternite about as long as 2n�. Joint 1 of metatarsi slightly shorterthan all following tarsomeres combined, andjo ints 2 and 3 each considerablyshorter than 4th. Body length: .09 mm. Diagnosis: Thisspecies differs fromCyanopenthemetallica (Champ.), comb. nov. (Plate 5: 10, 11 ), in thebroader pronotum(in metallica only twice as broad aslong) C. withfine and sparsepunctation (Plate 5: 9) (in C.metaUi ca, punctationof rather medium size,at least partly umbilicate,and spaces between punctures shining, rugose). Scutellum in C.metal lica of bronzed or metallic red-bronzed color, more or less evenly covered with dense yellow hairs, withoutdistinct impre ssion (Plate 5: 1 O).Antennomeres 8-10 in metallica less transverse (Plate 5: 11 ). C.
Genus Pen the Newman, 1838
Plate 5: 1-8 Type-species:He lops obliquatusFabricius , 1798 Thisgenus includes nine species distributed in China, theHimalaya, Japan, the Oriental Region, andNorthAmeri ca. Larvaas in Plate 16: 1-17. Probablymycetopha ges.
Penthe reitteri sp. nov.
Plate 5: 3, 4 Holotypemale, China, Sechuan,Nitou Tatsienlu, Em. Reitter. -Paratypes: 1 male, 1 female withthe same data. 33 Theholotype andone paratypeare kept in theNarodni Muzeum v Praze,Prague, whilethe other paratype in the ZoologicalMuseum of the StateUni versity of Moscow, Russia. Description. Body black, oblong-oval, at least elytra more or less dull; last antennomere yellowish. Upperside clothed with black or black-brown, somewhat raised,moderately dense and modestly elongate Pronotum at hindmargin before hairs. scutellumwith a transverse, U-shaped patch clothedwith dens e yellow hairs, in front of latter animpre ssed patch clothed with erect blackhairs . Scutellummedially with well-developedblack pu bescence surroundedwith denseyellow pu bescence (Plate 5: 3). Female antennomere 3 strongly elongate but a little shorterthan two sub sequent joints combined, 6th considerably shorterthan Sth; male antennomere 5 considerably broader and longer than 4th, 6th or 7th, latter considerably longer than 6th; male antennomeres 7-10 rathershort, subequal, 1.2-1.25 times longer than wide, densely clothedwith rather short decumbentpu bescence. Frontal widthbetween eyes 1.3-1.4 times less thantransverse ocular diamete r. Head posteriorlyand pronotum rather finely and densely punctured, punctures in fore part of head larger. Pronotum 2.1-2.2 timesas broad aslong, withsomewhat pointedly rounded and proj ecting posterior comers (Plate 5: 3). Pairedbasal impressionsof pronotum well developed, fovea-like,more or less longitudinal. Elytra twice aslong as maximum broad behind middle, each with 14 very fine rows of punctures (including presutural row but excluding a rough row on lateral border). Spaces between rows ea. 3-4 diameters of a puncturein row; spaces between rows withvery fine, sup erficialand dense punc tures, latter only relativelylittle smaller thanpunctures in rows (Plate 5: 3). Aedeagus asin Plate 5: 4. Body length: 10.5-12.0 mm. Diagnosis: Differs well fromcongeners by thefollowing combination of characters: coloration of pubescence of scutellum and of posterior part ofpronotum as well as veryfine rows of puncturesof elongated elytra.
Subfamily Hallomeninae Mulsant, 1856
Genera included: Hallomenus Panzer, 1794 andMy cetoma Dejean, 1834
Genus Hallomenus Panzer, 1794
Plate 6: 1-17; Plate 10: 1
Type-species: Hallomen ushum eralis Panzer, 1794 (= binotatus Quensel, 1790) (Plate 6: 1) 34
This genus is divided into two subgenera: Hallomen us s. str. (Plate6: 1-14; Plate 10: 1) with ten species distributed in Europe, the Caucasus, Siberia, the FarEast of Russia, China, Japan,Pakistan, andNorthAmerica, andXe uxes Champion, Kashmir 1889, stat. nov. (Plate 6: 15-17), knownfrom the southof the RussianFar East, Japan andNorthAmerica (including Mex ico). Xeuxes was described as a monotypic genus and included only X brevicollis Champion, 1889, from Mexico (Plate 6: 15, 16). While studying the world fauna of thisgroup, two more species have been found tobe actuallyreferred tothis subgen us: Hallomenus(Xe uxes) serricomis LeConte, 1877, from North America, and H. (X.) tokejii Nomura et Karo, 1958 (Plate 6: 17), distributed in the south of the Ru ssianFar Eastand in Japan.The latter species was earlier included in the monotypic subgenus Parahallom en usNomura et Kato, 1958. Examination of all three above species has allowed to establish absolute synonymy of Xe uxes Champion, 1889 and ParahallomenusNomura et Karo, 1958, syn. nov. We have examined the type of Xeuxes diversicomis Pie, 1930, described from Argentine, and found out that in reality it belongs to the subfamily Alleculinae of the fami lyTenebrionidae. Inaccordance with thecatalogue ofCsiki (1924), some species in fact belonging to other families or genera have also been included inHallomen us. Thus, Hallomenus pa/Jens Gyllenhal, 1817, described from Sierra Leone, is in reality referred to the familyScrap tiidae; we have studiedthe typesof speciesand designated the lectotype. this HaUomenusreticulatusMotschoulsky, 1872, described from Atlanta, Georgia, U.S.A. (theholotype of which also beenrestudied by us)actually belongs in thesubf amily has Alleculinae, family Tenebrionidae. Hallomenus fuscosuturalis Blatchley, 1913, described from Florida, U.S.A., is in reality another member of Scraptiidae. Unfortunately, the type of HaUomenus variega tus Motschoulsky, 1872, from the Caucasus, has not been recovered in the Motschoulsky Collection of the Moscow Museum but,based on theoriginal description, it is verylikely to represent the genus Orch esia Latr. (Melandryidae). Similarly, no typeof Hal/ omen us ?anas pioides Motschoulsky, 1845(a), described from Kamchatka, been elucidated in the Motschoulsky Collection. However, it has does contain a specimen which could have representedthis species, judged fromthe geographicallabel and regrettablyfar toobrief original description.Ye t it seems likely be non-type. Inreality it represents Orchesia Solsky, 1871 (Melandryidae). to tiisiformis Assoon asthere isno convincing evidence about the originalidentity ofMotschoulsky' s specimen, we conservethe latter nameas valid. Further, the species described asMy cetocharapuncti collis Blatchley, 1917, from North America, is actually referred to the genus Hallomen us,we have restudied the type of this species. Ha llomenus imlatus Kangas, 1959, described fromFinland (we have restudiedthe type aswell), belongs in factto the genusScrap tia Latr. (Scraptiidae). Based ona restudy of thetype, Ha Uomenusklapp erichi Pie, 1954, described from China, long become type-speciesof a separategenus, PseudoholostrophusNikitsky , has 35 1983. Finally, we consider Hallomenus punctulatus LeConte, 1866, described from NorthAmerica, as a junior synonym of H. bmotatus(Que nsel, 1790). Apparently, the development of HaUomen us species is mainly associated with Aphyllophorales fungi.
Ha llomenus (Hallomen us) chinensis sp. nov.
Plate 6: 8-1 1 Holotype male, China, Sechuan, Gongga Shan, above Camp 3, 3050 m, 22.VII. 1994,A. Smetanaleg. - Paratypemale, sameloca lity. The holotypeis kept in thecollection of theMuseum d' Histoire naturelle in Geneva, the paratype in theZoological Museum of theState University of Moscow, Russia. Description. Upperside and underside black-brown to black; coxae, trochanters and femora red-yellow, tibiae with tarsi brown or almost black; antennomeres 3-11 (excluding a weaklyclarified apex oflastjo int) black-brownto black; maxillarypalps, antennomeres 1-2 and anterior edge of labrum darkred-brown . Upperside covered withmore or less elongatehairs, latter lighter grayishon head andpronotum anddarker brownish-greyon elytra. Last joint of maxillarypalps ea. 1.8-1.9 times as long asbroad, considerablylonger at outer edge than at inner one, more or less roundly and obliquely truncate apically. more or less elongate, when stretchedback alongbody, reachingabout anterior Antennae 1/5 extent of elytra. Antennomere 1 distinctly longitudinal, 1.5-1.7 times as long as 2nd, latter short and about as long as broad; antennaljo int 3 elongate-triangular, ea. 2.2-2.3 times as long and 1.6-1.7 times as broad as2nd and 1.3-1.4 times as long as broad; antennomere4 ea. 1.15-1.25 times as short as 3rd, barelylongitudinal or about as long as broad; antennaljoints 5-10 about as long as broad; antennomere 11 elongate, ea. 1.6-1.7 times as long as 1 Othand 2.0 times aslong as broad, distinctly narrowed in apical partand conspicuously constricted(Plate 6: 9). Head ratherslig htly shining or almostdu ll, withdense, medium- sized, granulatepunctation, very finely shagreened, at most witha weakflattened impression level to posterior edge of eyes. Widthof frons between strongly sinuate eyes 1.15-1.2 times less thantransvers e ocular diameter. Pronotum generally more or less convex, 1.6-1.7 times as broad ashead witheyes andas its ownlength. Its sides beforeposterior roundly obtuse comers (approximately inposterior 1/4 length) at firstweakly broadenedand then stronglynarr owed towards anterior, fullyrounded comers (Plate 6: 8). Baseof pronotumdistinctly bisinuate, all edgesfully margined Disk of pronotummore or less shining, denswith e, rathermedium sized, granulate punctation (Plate 6: 8), withtraces offine shagreenin spaces at most. Distancebetween punctures in longitudinal direction ea. 0.5 diameterof a puncture. Paired basalimpre ssions of pronotum large and deep, but a smaller and shallower impression canbe situated in front of each. Medial surfaceof pronotum canbe weakly 36 transverselyflattened in posterior part,anterior part usually with a more or less distinct longitudinalimpre ssion. Scutellumabout as long asbroad, strongly narro wed posteriorly, more or less roundly truncate, with finepunctation and finely shagreened. Elytra oblong-oval, somewhat flattened,2.0 timesas long asgreatest width.Their surfacefinely anddensely punctured, more orless shining, with a conspicuous transverseimp ression in anteriorpart at suture. Each elytrumwith 5-7 weak striaebut without distinctlydeepened rows of punctures inside (Plate 6,8). Spaces between puncturesof elytrawith weak traces of shagreenat most, up to 0.5-1.0 diameter of a puncture in longitudinal direction. Prostemumbefore procoxae ea. 2.0 times as short as theirlongitudinal diameter. Trochantinof procoxae invisible. Prostemal process ratherbroad, margined on sides, more or less roundlybroadened posteriorly and inthis part ea. 2.0-2.2 timesas narrow as coxal transverse diameter. Mesostemal process narrow, margined, considerably narrower than prostemal one, almost reaching the posterior edge of mesocoxae. Metastemumfinely punctured in middle partand finely shagreened, ea. 2.0 times as long as 1 st visibleabdominal sternite. Metepisternaconspicuously narr owed posteriorly, ea. 3.5 times as long as broad, without well-defined triangular area in anteriorpart. Joint 1 ofprotarsi considerably longer than joints 2 and 3 combined, slightly longer than5th. Metatibiaeabout as long astarsi, their spurs short, of almost equallength, 5 .0 times as short as tarsomere 1, latter considerably longer thanjo ints 2 and 3 combined and ea. 2.2 times as long as 4th. lst visible abdominal sternite subequal to 2nd. Abdominalsternites with fine punctation and finely shagreened, more or less dull. Aedeagusas in Plate 6: 10, 11. In the holotype penis andparameres approximately in one plane (Plate 6: 10, 11) , while in the paratypethe penis is curved almost at right angleswith respect to theparamere s. Diagnosis: The species differs from other species of HaUomenus s. str. in the followingcombination of characters: granulatepunctation of pronotum, more or less uniform black-brownor black body with femora considerablylighter thantibiae (Plate 6: 8), andfairly strongly enlarged antennom eres.
Hallomen us (H. ) orientalis sp . nov.
Plate 6: 12-14 Holotype female, India, Kashmir, Gulmarg, VI.1931, Dr. M. Cameron leg. - Paratypes, 3 females, Pakistan, Hazara,Malkandi , 1500 m, 3.VI.1983, Besuchet & LObl leg. Theholotype is kept in thecollection of theNatural History Museum in London, two paratypes in theMuseum d'Histoire naturelle in Geneva, andone paratypein the Zoological Museumof the StateUni versity of Moscow, Russia. Description.Main color ofbody black-brownto dark red-brown; abdomen usually moreor less clarified; maxillarypalps, headgenerally or its anteriorpart only, pronotum 37
II
7 17
9
8
... . -:
6 5 19 18 20 21
Plate 6. General view and details of structure of Hallomeninae (all original). 1-6 - Hallomcnus (Ha llomenus)binotJJtus; 1 -H.(H.) nipponicus;S-11 -H.(H.) chinensis(type); 12-14 -H.(H.) oricntalis; 15-16 -H.(Xc uxcs) brcvicollis (type); 17 -H.(X.) tokcjii; 18 - My cctoma suturalis; 19 - M ussuricnsis Nikitsky, 1985; 20, 21 - M af1inis Nikitsky, 1985. 1, 8, 18 - general view; 2 - maxillary palp; 3 - metendosternite; 4 - hind leg;5 - aedeagus (dorsal view); 6 - same(lateral view); 7, 9, 14, 15, 17, 19 - antenna; 10, 20 - tegmen (dorsal view); 11, 21 -penis; 12, 16 - pronotum; 13 - elytrum. 38 (except fora transverse obscure dark fas cia usually divided in middle partinto two spots by a more or less light longitudinal area), antennae and legs red-brown or red yellow; largehumeral spot on each elytru.mred-yellow and slightlynot reaching the suture, but reaching the lateral edge (Plate 6: 13). Antennomeres 3-10 more often strongly more rarelydark red-brown. Upperside coveredwith elongate in:fu.scate, hairs, more or less decumbentor stronglyinclined, lighton disk ofpronotum andinfuscate againstblack-brown or almost black backgroundof elytra. Antennomere I longitudinal, ea. 1.4-1.6 timesas long as2nd; latter short, about as long as broad; antenna!jo int 3 elongate-triangular, twice aslong as 2nd and 1.3-1.4 timesas long as bothwidth and antennomere 4; antenna!jo ints 4-10 more or less equal insize, antenna!jo ints 8-10 ea. 1.1-1.2 timesas broad as long; 11th more or less oblong oval, ea. 1.4-1.5 times as long asboth wide and lOth(Plate 6: 14). Head with dense, ratherfine, granulate punctation; distancebetween deeplyemarginate eyes ea. 1.4-1.5 timessmaller than ocular transversediameter. Pronotum more or less shining, stronglytransverse, ea. 1.7 times asbroad asboth long andhead witheyes ; pronotum broadest in posterior part,whence its edges fairly stronglyand roundly narro wedanteriorly. Anteriorcomers of pronotumentirely rounded, posteriorones also rounded butless strongly so. Baseof pronotumshall owly bisinuate. Granulatepunctation of pronotummoderately large, very dense,punctures contiguous or almostso, disk of pronotum somewhatflattened in middle part. Scutellum roundly triangular, finelypunctured, very weakly transverse. Elytra oblong-oval, 2.1-2.3 times aslong asmaximum width, more or lessshining, somewhat flattenedon disk, at base only barelybroader than pronotum, with dense, fine,rugose punctation(considerably finer than on pronotum) andat most withvery weak traces of two striaein anterior part of elytra. Prostemal process considerably broadened androunded caudally, its widthbefore broadestplace ea. 3.5 timesas narro w ascoxal transverse diameter. Mesostemalprocess almostreaching the posterior edge of mesocoxae. Sides of metastemumand its epistema withdense and rough puncta tion. Spursof metatibiae ea. 5.0 times asshort asjo int 1 of metatarsus, ea. 1.2 timesas short as followingjoints combined; joint 2 considerably latter longer than3rd andsub equalto 4th. Abdominal stemites finelyand densely punctured, shining. Body length: 4.4-5.0 mm. Diagnosis: This formdiff ers fromother species ofHaUomenuss. str. in combining the following characters:pronotum with granulate punctation, reddis h-yellow except formedial areainfuscate like a transverse, dark, obscure fascia usually divided into two spotsby a lighterlongitudinal patch in middlepart (Plate 6: 12); elytrafairly strongly elongate (ea. 2.1-2.2 times aslong asmaximum width) , theirhumeral spots yellow brown,fai rly sharplydelimited from remaining surface of elytra(Plate 6: 13), far from reaching the suture but enveloping humeral tubercle; medianpart of elytrawith two weakstriae at most. 39 Key to species of based on larvae Ha llomenus Plate 18: 1-10 l. Urogomphimore or less straightin lateral view , retlexeddentiform atapex and at most only very slightly curved (Plate 18: 9, 10). Bodylength: 5. 0-6.Smm. Europe,
Siberia ...... (Illiger, 1807) Caucasus,(?) H. axillaris -. Urogomphi rather evenly andmore or less strongly curved all along in lateral view, curvedf orward atapex (Plate 18: 7). 6.5-9 .0 mm.Europe , Siberia Caucasus, ...... bino'fatus (Quensel, 1790) H.
Genus Dejean, 1834 My cetoma
Plate 6: 18-21 'fype-species: Dryops Panzer, 1797 suturalis This Palearcticgenus isrepresented in the world fauna by fourspecies, of which one isdistributed in Europe andthe one morein the southof the Khabarovsk Caucasus, andMaritime provinces (Nikitsky, 1985), further one inJapan and Island, Kunashir while the fourthin Japan. Theirdevelopment ismainly associated with fungi of the genus/sclmoderma. Larvaas inPlate 17: 1-11.
Subfamily Eustrophinae Gistel, 1856
subfamilyis dividedinto two tribes: Holostrophinitrib . nov. andEustrophini This Gistel. The former includes two genera: Holostrophus Hom, 1888 and Pseudoholostroph us Nikitsky, 1983. The latter tribe is composedof three genera: EustrophusIlliger , 1802, Syns troph usSeidlitz, 1898, andEustrophopsis Champion, 1889.
Tribe Holostrophini, tribus nov.
Genera included: Ps eudobolostropbusNikitsky Ho and lostropbusHom Representatives of differfrom the Eustrophini in the meten dostemitewith thistn"be well-developed plates (Plate9: 12) aswell asin the morecomplex lateral directed laterad structureof the aedeagus:basal part of tegmenstrongly elongate,p ara.meres likewell developed andstrongly separated lobes, apicesoflatt er oftensclerotized more weakly thanbasal part. Penis stronglyelonga te, witha spiniform structurein its anteriorpart 40 and along rod-shapedsclerotized process ventral on side. Inaddi tion, simplemetatibiae withouttransverse of rows spinescombined with confused punctation of theelytra are not characteristic of the true Eustrophini. The larval labium which is divided into a prementum, a mentumand asubmentum. more or less distinctly separated from the gulain Holostropbus, aswell as thestructure of themandibl es, the well-developed epicranialstem and the presence of a toothat the inneredge of theurogomphi are not characteristicat least of the larvae of Eustropbopsis (= Eustrophin us) , i.e. the only genus ofEustrophiniin which thelarva is suffi cientlywell-describe d.
Genus Nikitsky, 1983 Pseudobolostropb us
Type-species: HsllomenusJdap pericbi 1954 (Plate 7: 3-8) Pie, The genus been established based on a study of the type of Ha llom en us has Jdappericb iPic, 1954, a speciesdescribed from Kuatun, Fukien,Chi na. It differsfrom HolostropbusHom, 1888 in the smallerand moreweakly emarginate eyes, withthe betweenthem being considerablygreater than a transverse oculardiameter distance (in Holostropbus, less thana transverse oculardiameter), as well asin the prostemal processusually faintly to not projectingbehind the posterior edge of theprocoxae and largely entirelymargined on sides (inHo lostropb us, prostemal process considerably longer andnot rarelyvery strongly protruding behind posterior edge of procoxae) . Pseudobolostropb usishere considered to encompass four species,including also Pseudobolostropb us cbinensis sp. nov. from Sechuan, China (Plate 7: 1, 2), Pseudobolostropb usimpressi collis(J...A£on.te, 1874), comb.nov. exHo lostropb us(Plate 7: 9-11), andPseudob olostropb usdiscolor (Horn, 1888), comb. nov. ex Ho lostropb us (Plate7: 12-15). Bothlatter speciesare NorthAmerican, while discolor (we have seen only a single specimen) is promoted to type-species of a separate subgenus, Holostrophinussubgen.nov. Thissubgen us ersdiff from Ps eudob olostropb us s. str. in thedouble punctation of thepronotum. (Plate7: 12) and the weaklyimpre ssed pairof basalrather nonstrok.e-shaped impr essionsof thepronotum. It seems noteworthy thatspecies of Ps eudob olostropb us display the elytraeither one-colorbut not blackor witha lighthumeral spot(Plate 8: 1) only, or with a clarified diffusedtransverse fascia inbasal part (Plate 7: 12). A more clearlyevident reddish yellow or red spotty pattern of the elytra is characteristic of Holostropb us, not Pseudobolostropb us. The larvais unknownto us.
Key to species of based on imago Pseudobolostropb us
1. Pronotumwith evident double punctation, withrather dense and largepunctures interspacedby smallerone s (Plate 7: 12). Pair of basalimpressions of pronotum 41
1 2 GD 11
9 13 8 10
12
15
5
6 7 � 3 4
Plate 7. General viewand details of structure of Pscudoh olostrophus spp. original). 1, 2 - (all Pscudoholostrophus(Ps cudoholostrophus) ch incnsis (type); 3-8 -P.(P.) Jdappcrichi (type); 9-11 - P. (P.) imprcssicollis; 12-15 -P.(Ho lostrophinus) disco/or: 1, 14 -head (frontalview); 2, 11, 15 -prostermnn (ventral view); 3 - general view;4 - maxillarypalp; 5 - sternum (ventral view); 6-head andprostemum
(lateral view, schematic); 7, 10, 13 - antenna; 8 - penis (dorsal view); 9 - head, pronotum and elytra;
12 - pronotumand elytra; a - mesepistemum; b - mesepimerum; c - triangular area of anteriorpart of metepistemum; d - metepistemum. 42 rathersh allow, rather nons1rokein shape.Prosternal p rocess considerablybroadened posteriorly, slightlybut distinctly surpassingposterior edge ofprocoxae, bent under mesosternumbut its vecytip weakly elevated (Plate7: 15) ...... Subgenus Holostrophinus subgen. nov. Body black-brown to black; maxillary palps, basal 4-5 antennomeres and antennomere 11, aswell aslegs red-brown; antennomeres 6-10 usually infuscate, underside clarified at least partly, as arule . Basal part of elytrawith a diffused, transverse,red-brown or red fasciaturning behind into darkerbackground of elytra withoutany sharpbound aries.Upperside coveredwith somewhat raised, elongate,
more or less infuscate hairs. Body length: 4.6-5.5 mm. NorthA.merica...... P. (I/.}dis color (llom) -. Pronotum with simple, more or less uniform punctation(Plate 8: 1 ). Pair of basal impressionsof pronotumusually distinctly stroke-like or in any casedeeper (Plate 8: 1). Elytra one-color (Plate 7: 3, 9) or with a light, rather contrasting humeral spot (Plate 8: 1) ...... Subgenus Pseudoholostrophus s. str .... 2 2. Bodyblack, each elytrumwith a red-yellowhumeral spot(Plate 8: 1 ); antennomeres 1-3 or 1-4, labrumand maxillary palps dark red-brown; apex of terminal antenna! joint andtarsi somewhat clarified.Elytra at leastagainst dark background covered with dense , latterblack-brown, slightly raisedand somewhatdarker thatthose hairs on pronotum. Pronotum rather weakly shining or more or less dull, at least its middlepart with moderately dense, rathermedium- sizedand not deep punctation (distance between punctures subequal to or a little greater than their diameter).
Body length:4.5 mm. China ...... (P.) chinensis sp. nov. P. -. Body one-color, red-brownto brown, not rarelyred tingeon upperside. Pronotum withfine or roughbut densepunctation (distance between punctures considerably less thantheir diameter) ...... 3 3. Pronotum and elytrawith dense and rough punctation, shining, not shagreened (Plate 7: 3). Stroke-like impressions of pronotum short and occupying ea. 1/5 pronotal length (Plate 7: 3). Upperside covered with dense, semi-erect, reddish brownhairs. Pronotum 1.6-1.7 times asbroad aslong. Body more or less convex, darkred-brown to almostbrown; disk ofpronotum more or less infuscate,basal 2-
3 antennomeres andapex ofjoi nt 11 clarified. Body length: 5.1-5.4 mm. China ...... P. (P.) klapp erichi (Pie) -. Upperside densely and finely punctured, weakly shining or dull. Stroke-like impressions ofpronot um fairlylong and occupying about or slightly lessthan 1/3 pronotal length (Plate 7: 9). Upperside covered withrelatively short, grayish or yellowish-gray, decumbent hairs. Pronotum less transverse, ea. 1.4-1.5 times as broad aslong. Body oblong-oval,red-brown, generally moreor less convex. Body
length:5- 7 mm. NorthAmerica ...... P. (P.) impressicollis (Leconte) 43 Pseudoholostrophus (Holostrophin us subgen. nov.) discolor (Hom, 1888), comb. nov.
Plate 7: 12-15 Material: 1 ex., North.America. Body black-brown to black; maxillary palps, basal 4-5 antennomeres and antennomere 11, as well as legs red-brown; antennomeres 6-10 usually infuscate, undersideclarified at leastpartly, as a rule. partBasal of elytrawith a diffused,transverse, red-brownor red fascia turningbehind into darkerbackground of elytrawithout any sharp boundaries. Upperside covered with somewhat raised, elongate, more or less infuscate hairs. Apical joint of maxillarypalps about as long asbroad, obliquelytruncate at apex. Antennae comparatively shortand usually slightlynot reachingthe posterior edge of pronotum, theirfour apical joints broadened considerably morestrongly than preceding ones; antenna!joi nt 1 ratherlarge, ea. 1.7 times as long asbroad; joint 2 ea. 1.6 times as shortas 1st and ea. 1.25 times aslong asbroad; joint 3 elongate-triangular, ea. 1.4 timesas longas 2nd and ea. 1.7 times aslong as its ownwidth; joi nt 4 about aslong as 3rd and 1.4-1 .45 times as long as broad; joint 5alsolongitudinal, only veryslightly shorterthan 4th; joint 6 ea. 1.1-1.15 times asshort as 5thand longer thanwide; joint 7 ea. 1.15 times as short as and very slightly broader than 6th, ea. 1.25-1.3 times as broad as long; joint 8 ea. 1.2 times asbroad asand subequal in lengthto 7th, 1.6 times asbroad as long; joint 9 slightly shorter, 1.2 times as broadas 8thand ea. 2 times as broad as long; joint 10 veryslightly shorter andbroader than and2.3- 2.4 timesas 9th broad aslong; joint 11 oblong-oval,subequal in lengthto two preceding onescom bined, 2.7 times aslong as lOthand 1.2 times aslong asbroad (Plate 7: 13). Head withdense and fine punctation (distance between punctures considerably smaller than their diameter), distinctly flattenedin medial partbehind eyes; distancebetween relatively small andweakly sinuate eyes ea. 1.6 as greatas ocular transverse diameter. Pronotum convex, ea. 2.5 timesas broad ashead with eyes and 1.45 times asbroad as long, margined along all edges, its base shallowly bisinuate, more or less straight truncatein middle part.Pronotum broadest at posteriorea. 1/4 extent, whence its sides strongly and roundly narrowed anteriorly, while posterior comers more or less rectangularly rounded. Disk of pronotum more or less dull or weakly shining, with traces of very fineshagreen andwith double punctation of coarse, moderatelydense puncturessomewhat elongate in longitudinal direction(distance between them inmiddle partof disk 0.3-1 diameterof a puncture) and finepunctures in between (2-3 times smallerthan coarse one s). Scutellum more or less roundlyquadrangular, dull, ea. 1.3 timesas broad as long, withde nse, fine punctation andfinely shagreened. Elytra oblong-oval, more or less convex, at humeri aboutas broad aspronotum at base, 1.7 times aslong asbroad, their 44 disk with coarsepunctation and partly with tracesof fineshagreen, thus making elytra look like more or less weaklyshining oralmost dull. Distance between punctures on disk of elytra (subequal to those on pronotum) in transverse direction up to 0.3-0.7 diameterof a puncture and, in longitudinal direction, subequal to theirdiameter (Plate 7: 12). Presutural striawell-developed. Sides of prostemum also with moderately dense double punctation, but lacking dense rugose sculpture. Prostemum before procoxae ea. 2.2 times as short as their longitudinaldiame ter. Prostemal process in its broadened part1. 7- 1.8 times as narrow as coxaltransverse diameter , considerablybroadened posteriorly, slightlybut distinctly suipassingposterior edge of procoxae, bent undermesoste mum but itsvery tip weakly elevated (Plate 7: 15). Mesosternalprocess without distinct impression inmedial part, reaching ea. posterior 114-115 length of mesocoxae. Sides of metastemum and its epistema also with double punctation. Metepistema distinctlynarr owed posteriorly, ea. 2.8 times as long as broad. Metastemum considerably longer than lst visible abdominal stemite, latter by far more finelypunctured. Spurs of metatibiae subequal in length,ea. 3.5-3.8 times as short as tarsomere 1, latter ea. 1.5 times as long as joints 2 and 3 combined, joint 2 considerably shorterthan 4th. 1 st visible abdominal stemite subequal to 2nd. Body length: 4.6-5.5 mm. NorthAmerica.
Pseudoholostrophus (P seudoholostrophus) chinensis sp. nov.
Plate 7: 1, 2; Plate 8: 1, 2 Holotype female, China, Sechuan, Gongga Shan, above Camp 3, 3050 m, 22. VII.1994, A. Smetana. Thetype is kept in thecolle ction of theMuseum d' Histoire naturellein Geneva. Description. Bodyblack, each elytrumwith a red-yellow humeral spot (Plate 8: l); antennomeres 1-3 or 1-4, labru.m maxillaryand palps darkred-bro wn;apex of terminal antennaljo int and tarsi somewhat clarified. Elytra at least against dark background covered withdens e hairs, latter black-brown,slig htly raised andso mewhat darkerthat thoseon pronotum. Pronotumrather weakly shining or more or less dull. Joint 4 of maxillarypalps ea. 1.6 timesas long as broadand ea. 2.2times as long as 3rd. Antennae rather short and, when stretched back along body, barely surpassing base of elytra.Antennomere 1 stronglydeveloped, longitudinal, considerably longer than2nd, latter ea. 1.6 timesas long as broad; joint 3 elongate-triangular, ea. 1.1-1.2 timesas long as 2nd and 1.8 times aslong asbr oad;jo int 4 also distinctlylongitudinal, ea. 1.3 times as short as 3rd; joint 5 ea. 1.3-1.4 times broader andvery slightly longer than4th, barely longitudinal; jo ints 6-10 more or less graduallybroadened, transverse, almostequal in length;joi nt 6 ea. 1.25 times asbroad as long; joint 10 ea. 1.8 times as broad aslong; joint 11 ea. 2.5 times longer thanOth 1 and ea. 1.2-1.3 times as long as broad(Plate 8: 2). Head more or less dull, weakly impressed medially behind eyes, 45 withde nse, rathernot coarsepunctation and markedly, finely, transversely shagreened. Width offrons between theeyes 1.25 times greaterthan oculartransverse diameter (Plate 7: 1). Pronotum stronglytransvers e, twice as broad ashead with eyes and ea.1. 6 times asbroad aslong, ratherweakly shining or more or less dull, at leastits middle partwith moderately dense, rather medium-sized and not deep punctation (distance between punctures subequal to or a little greaterthan their diameter). Pronotum broadest at base,whence anteriadat firstmore or less parallel-sided(about posterior 113 extent) and then strongly and roundly narrowed. Base of pronotum more or less trisinuate, basalimp ressions stronglydevelop ed, stroke-like andextending up to ea. 1/3 extent. Posterior comers ofpronotum roundly rectangular, anterior ones entirely rounded. Middle of pronotum weakly transversely impressed against a more or less convex general background. Scutellumfinely puncturedand shagreened, more or less straighttruncate at posterior edge(only barelyrounded) . Elytraoblong-oval, middlepart of disksome what flattened, at humeri barelybroader thanpronotum at base, ea. 1.85 times as long as maximum width. Elytrashining, virtuallynot shagreened; punctationdens e, transverselyrugose, considerably denser and markedly coarser and deeper than on pronotum; average distancebetween punctures in transverse directionconsiderably less thantheir diame ter, in longitudinal direction a little greater than or subequal to diameter of a puncture. Apices of elytramore or less broadly rounded. Prostemumbefore procoxae ea. 1.6 times asshort as coxal longitudinal diameter. Prostemalpr ocess rather finelypunc turedand finelytransversely shagreened, markedly broadened towardap ex, margined along sides, virtuallynot projectingbehind posterior edge of procoxae, in its broadest partea. 2.3-2.5 times as narrow as coxal transverse diameter. Mesostemal process narrower thanprostemal one, margined, reachingthe posterior 1/3 extent of mesocoxae and inits posterior part ea. 4.5 times as narrow as coxaltransverse diameter . Longitudinal metastemal suture ea. 1.3 timesshorter than metastemum(measuring its diameterfrom meso- to metacoxae) . Sides of meta.sternum withmoderately dense, coarsepunctation. Metepistema(regar dless of a well-defined, triangular, somewhat elevated areaanteriorly) ea. 3 times aslong as broad, distinctly narrowed posteriorly. Most part of sternum and its epistema evidently, finely and transversely shagreened. Shagreen of abdomen also well-developed. Metatibiae sub equal in lengthto metatarsi,their spurs ea. 4 times as short as tarsomere 1, latter barelyshorter thanall following tarsomere s combined; tarsomere2 ea. 1.4-1.5 timesas long as 3rd andbarely longer than4th. Body length: 4.5 mm.
Pseudoholostroph us (P. ) klapp erichi (Pie, 1954)
Plate 7: 3-8 Lectotype male (designatedherein), China, Kuatun, Fukien, 22.X.1946, Tshung Sen leg. - Paralectotype, same data (see Nikitsky, 1983). 46
The lectotype ishoused in the Staatliches Museum fiirNaturkunde in Stuttgart, the paralectotypein theMuseum national d'Histoire naturellein Paris. Description (see also Pie, 1954a and Nikitsky, 1983). Body more or less convex, darkred-brown to almostbrown; disk of pronotum more or less infuscate, basal2-3 antennomeres and apex of joint 11 clarified. Upperside covered with dense, semi erect, reddish-brownhairs . Antennae relatively short and, when stretched back along body, only relatively slightly surpassingbase of elytra.Antennomere 1 fairly strongly developed, longitudinal, ea. 1.6- 1.8 times as long as 2nd; latter ea. 1.25 times as long as broad and 1.2 times as short as 3rd; latter 1.6 timesas long as broad and ea. 1.2 times longer thana distinctly longitudinal antennomere 4; antennomere 5 ea. 1.25 times as long as broad and I.I times broader than 4th; antennomeres 6- 10 more or less gradually broadened; antennomere 6 about aslong as broad, I Othea. 1.7 times asbroad as long; antennomere 11 oblong-oval, ea. 2.3-2.4 times longer than IOthand 1.4 times as long as broad (Plate 7: 7). Head more or less shining, densely, relatively finelypunctured. Distance betweenrather weakly sinuate eyes ea. 1.3 timesgreater than ocular transverse diameter. Stroke-like impressions of pronotum short andoccupying ea. 1/5 pronotal extent (Plate 7: 3 ). Pronotum 1.6-1.7 times as broad as long, stronglytransvers e, about twice asbroad as head witheyes, its basetrisinuate andposterior comers more or less obtusely rounded. Pronotum and elytrawith dens e and rough punctation, shining,not shagreened (Plate 7: 3). Medial part of pronotum before scutellum andbefore middle withweak impressions. Punctures on pronotum coarse, considerably larger than on head,very dense, spaces between them likevery thin wrinkles, 2-3 timesnarr ower thandiameter of a puncture. Scutellum transverse, ea. 1.5 times as broad as long, rather finely and densely punctured, eithermore or less straighttruncate at posterioredge or weakly rounded. Elytra oblong-oval, at humeri subequal in width to pronotum at base, 1.93 times as long as broad, withdense andrough, transverselyrugose punctati on, puncturesbei ng about same size as on pronotum; average distance between punctures considerably less thantheir diamet er. Prostemumbefore procoxae 1.7- 1.8 times asshort ascoxal longitudinal diameter. Prostemal process distinctly broadened posteriorly, almost not projecting behind posterior edge of coxae, without sharp margin along sides of apical part, ea. 1.8-2.0 timesas narrow ascoxal transvers e diameter (Plate 7: 5). Mesostemalpro cess more or less margined, narrowed posteriorly, in its posterior partea. 3.4-3.8 times as narrow as coxal transverse diameter. Longitudinal metastemal suture ea. 1.5 times shorter than metastemum. Sides of metastemum withvery rough andmoderately dense punctation anddistinctly but finely shagreened.Metepisterna also with dense punctation, but latter more finethan that of metastemum, also distinctly and finely shagreened; theirlength (regardless of a 1riangulat; well-isolatedarea anteriorly) ea. 2.8 timesgreater than width. Metastemum between meso- and metacoxae ea. 1.6 times as long as st1 visible 47 abdominal stemite. Metatibiae a little shorter thanmetatars i; tarsomere 1 subequalin lengthto all followingjoints combined; tarsomere2 ea. 1.5 times aslong as3r� latter twice shorter than4th. Penis asin Plate 7: 8. Body length: 5.1-5.4 mm. China.
Pseudoholostroph us (P. ) impressicollis (LeConte, 1874), comb. nov.
Plate 7: 9-11 Material: 7 males and females, North.America. Description. Body oblong-oval, red-brown, generally more or less convex. Upperside covered withrelatively short, grayish or yellowish-gray, decumbenthairs. Upperside densely, finelypuncture� weaklyshining ordu ll. Joint 4 of maxillarypalps elongate, ea. 1.5-1. 7 times as long asbroad and2.3-2.5 times aslong as2n� obliquely truncate at apex. Antennae rather short andusually not or barelysurpassing posterior edge of pronotum; antennomere 1 largeand longitudinal, ea. 1.5 times longer than2n� latter ea. 1.4-1.6 times as long aswide ; joint 3 strongly elongate, 1.2-1.35 times longer than 2nd and 1.6-2.0 times as long as broad; joint 4 subequal in length to 2nd and 1.35-1.6 times as long as broad; joint 5 subequal in lengthand width to 4th; jo int 6 verysligh tly shorterand broader than5th, ea. 1.1 times aslong asbroad; joint 7 subequal in lengthto andvery slightly broader than6th, about as long asbroad (barelylongitudinal at most); joint 8 ea. 1.1 timesas broad aslong and 7th;joi nt 9 ea. 1.3-1.4 times asbroad as long and 8th; joint 10 a little shorterand 1.1- 1.15 times broader than ea. 1.7-1.8 times as broad aslong; joint 11 more or less 9th, oval, ea. 1.8-2.0 times longer than lOthand 1.1-1.2 times as long asbroad (Plate 7: 10). Head finely andrather densely punctured and finely shagreene� dull. Width of fronsbetween weaklysinuate eyes 1.5-1.6 times thantransverse ocular greater diameter. Pronotum ea. 1.4-1.5 times as broad as long, about twice asbroad as head with eyes, its surface rather weakly convex, usuallywithout distinct impressions in medial part, withtraces of fine shagreenand dens e finepunctation, average distancebetween puncturesbeing considerablyless thantheir diameter. Baseof pronotumvery shall owly bi- or trisinuate, stroke-likeimpre ssionsfa irly long and occupying about 1/3 or slightly less pronotal extent (Plate7: 9). Pronotumbroade st at posterior 1/3 or 1/4 extent, whence its sides narrowed weaklyposteriorly and fairly strongly androundly anteriorly. Hind comers of pronotum obtusely roundedbut more distinct thanentirely roundedanterior ones. Scutellumtransv erse(ea. 1.8-2.0 times as broad aslong) , roundlytriangular, with veryfine punctation and traces of fine shagreen. Elytra moreor less convex, ea. 1.7 5- 1.85 times as long as maximum width, at humeri barely broader than pronotum at 48 base,at firstweakly broadened posteriorly and then roundly narrowed toward apex. Surface of elytrawith dens e, very fine,transversely rugose punctation (on the average finerthan on pronotum) andwith traces of fine shagreen. Distance between punctures intransverse direction considerablyless than,while in longitudinaldirection sub equal to,their diameter. Prostemum before procoxae roughly punctured and finely shagreened, more or less dull, beforeprocoxae ea. 1.1-1.2 times as short as coxal longitudinal diameter. Prostemalprocess ratherfinely punctured and distinctly shagreened, entirelymargined, surpassing posterior edge of anterior coxal cavities not more than 1/6 their length, distinctly broadened posteriorly, rounded at posterior edge, in broadest part ea. 1.8 timesas narrow as coxal transversediameter. Mesostemal process margined,roughly and densely punctured, shining, reaching approximately to posterior 1/3 extent of mesocoxae,without excavation for accommodationof prosternalpr ocess. Metastemum along midline subequal to of 1 st and 2nd visible abdominal sternites combined. Longitudinal metastemal suture anteriorly only slightly surpassing mid.length of metastemum. Sides of metastemum with rough, moderately dense punctation and distinctly shagreened. Metepistemawith finerpunctation than on sidesof metastemum, their length ea. 2.5 times exceeding the width. Metatibiae about aslong asmeta tarsi. Spursof metatibiae subequalin length, ea. 4.5-5.0 timesas short as tarsomere 1, latter onlybarely shorterthan all following tarsomeres combined; tarsomere2 ea. 1.7 times aslong as3rd andsubequal to 4th. Pr ocessof 1 st visible abdominalstemite separating metacoxae narrowly triangular. 1 st visible abdominalstemite a littlelonger than 2nd. Abdomen densely, veryfinely punctate and finelyshagre ened. Lastvisible abdominal stemite rounded at apex. Body length: 5-7 mm. North.America.
Genus Holostrophus Horn, 1888
Plate 8: 3-9; Plate 9: 1-17 Type-species:Eus troph us bifasciatus Say, 1824 (firstdesignated herewith) This genus encompasses 17 speciesin theworld fauna. These are distributed in the south of the RussianFar East, in Korea, China, Japan, the Oriental Region andthe Nearctic. Larvaas in Plate 19: 1-11; Plate 20: 1-6. Development basicallyin arborealfungi.
Holostroph us minimus sp. nov.
Plate 8: 7-9 Holotype female, Philippines, S. Luzon, Quezon N. P. (Lucena), 250 m, 8-10.1.1991, Bolm leg. 49
; , I , � , f', , " I I : ' 1, . I 4 . I�·' :�• l � �!{�:} •./,/..
5
9
8 2
7 3
Plate 8. General view and detailsof structure of Pseudoholostroph usand Holostroph us spp. (all original). 1, 2 - Pseudoholostrophus ch inensis; 3-6 - Holostroph usdiversef asciatus; 1-9 - H.minimus.
l, 3, 7 - 2, 8 - - 9 - 6 - general view; antenna; 4 head (frontal view); S, prostemum; aedeagus. 50 Thetype is kept in thecollection ofthe Naturhi storischesMuseum in Basel. Description. Red-brown, pronotumwith vague infuscate spots in anteriorpart on each side of middle andof the very posterior edge. Antennomeres 6-10 infuscate, black-brown; joint 11 red-yellow. Elytra black, each at base with a transverse red yellow fascia somewhat sinuateat posterior edge andextending forwardalong suture up to scutellum; a not very largered-yellow spot at sutureclose to middle; anda red yellow transversespot in apical 1/3 far fromreaching the suturebut reachingthe lateral edge;apex of elytrain presuturalarea somewhat clarified(Plate 8: 7). Uppersidecovered with more or less decumbent hairs, latter black indark areas andlight in red-yellow ones. Antennaerelatively short, antennomere 1 considerably longer than2nd, latter ea. 1.3 timesas long asbroad, considerablybroader andea. 1.15 times shorterthan elong ate triangular3rd one, latter ea. 1.6 times as long as broad; joint 4 ea.· 1.3 times aslong as broad and 1.25 times as short as 3rd; joint 5 ea. 1.3-1.4 times broader than4th and barely asbroad as long;jo ints 6-10 more or less graduallybroadened, similarin length; joint 10 ea. 1.8 times asbroad aslong ; joint 11 oblong-oval, ea. 2.5 times longer than lOthand 1.6 times as long as broad (Plate 8: 8). Head with rather dense and fine punctation and finely, transversely shagreened. Width of frons between eyes ea. 1.5 timesless thanocular transverse diameter. Pronotum moderately transverse, ea. 2.8 times broader thanhead with eyes andea. 1.4-1.5 times as broad aslong. Pronotal surface withrather dense and fine punctation (average distancebetween puncturesin middle partof disk up to 0.6-1.0 diameterof a puncture)and finely, transverselyshagreened, more or less shining. Pairof basalstroke like impressions of pronotum short, ea. 12 times asshort aspronotum . Pronotal base marginedonly in medial part, veryslig htly outreachingthe pairedbasal impressions; sides ofpronotum andits anterior edge entirelymar gined. Disk of pronotumdistinctly convex. Scutellum finelypunctured and distinctlyshagreened, transverse-quadrangular, ea. 1.8 times asbroad aslong. Elytraoblong-oval, ea. 1.65-1.7 times aslong as broad, at baseabout similarin widthto pronotum, shining(not shagreened), ratherfinely, densely, transverselyand rugosely punctured (distance between punctures in transverse direction apparently less thantheir diameter) , punctationbeing markedlydenser and verysli ghtly finerthan on pronotum. Prostemum before procoxae ea. 3.5 times as short astheir longitudinal diameter. Prostemalprocess ea 2.2 timesas ownarr ascoxal transv ersedi ameter, relativelysligh tly surpassingprocoxae (Plate 8: 9) andfai rly strongly bent under sternum; its partroundly projectingbeyond procoxae ea. 2.5-2.6 timesas short asmaximum width of process. Sides of prostemumwith fine shallowpunctation (without rough rugosity) . Impression on mesostemum receiving the prostemum weakly developed, short and narrow, representedonly in the very anteriorpart of mesostemum. Sides of metastemumwith punctation more sparse andrough thanits epistema,both sides and epistema distinctly 51 transverselyshagr eened. Spursof metatibiae subequalin length, ea. 2.7 timesshorter thantarsomere I. Joint 1 of metatarsus ea. 1.2-1.3 times aslong as2nd and3rd ones combined; joint 2 a littlelonger than3rd anddistinctly shorter than 4th. Body length:3.5 mm. Diagnosis: Thisspecies differs from congeners in combining such characters as coloration of the upperside, shortprosternal process, more or less smooth(without roughrugose punctation) sides of prostemum (hypomera), etc.
Holostroph us similis sp. nov.
Plate 9: 3-10 Holotype male, Malaysia, Sarawak, 4th Division Mulu nr. BaseCamp, Gn. NP, 50-100 m, Malaisetrap . - Paratype, 1ex., Sarawak, 4th Division Mulu nr. Gn. NP. , Camp, 150-200 Malaise VI.1978. m, trap. The typesare kept in theNatural History Museum in London . Description. Underside, legs, maxillarypal ps, head, antennomeres 1-5 andmain colorationof pronotum vagueinfuscate p on pronotumratherT-shaped red-brown; attern withtwo infuscate areas on each side ofbasalpart. Terminal antennaljoint veryweakly infuscatebasall y, more or less red or red-brownthere, yellowish apic ally. Elytra black, each with fourred or red-yellow spotsand a clarifiedapex; front spotof elytrasituated nearscutellum along suture; secondone larger, moreor less reachingthe lateral edge, lyingouts ide of front spot;medial spot situatednear middle of suture of each elytrum; andsub-a picalspot transverse, considerablyfar fromreaching the suture, situatedin apical 113 of each elytrum(Plate 9: 3). Uppersidecovered withmore or less elongate pubescence, only barely inclined,darker in black areas of elytra. Terminaljo int of maxillarypalps obliquely truncate atapex, subequal in lengthand width.Antennaerelatively short andusuall y, when stretchedback alongbody, slightly not reaching the posterior edge of pronotum. Antennomere 1 large andlongitudinal, considerably larger than2nd, latter 1.3-1.5 times as long asbroad; joint 3 elongate triangular, ea. 1.7- 1.8 times aslong asbroad and 1.3-1.4 timeslonger than 2nd; joints 4 and 5 a little shorter than 3rd,sub equal in length, but 5th slightlybroader than4th, about as long as broad; joints 6-10 more or less graduallybroadened; joint 6 ea. 1.1 times broader than 5th and ea. 1.2-1.3 times as broad as long; joint 10 subequal in length to 9th, 1.7-1.8 times broader than long; joint 11 elongate-oval, 2.1-2.3 times longerthan 1 andea. 1.4-1.6 times longer thanwide (Plate 9: 4). Head withdens e, Oth fairly finepunctation and finely shagreened, more or less dull. Width offrons between eyes 1.5-1.6 times as less thanocular transverse diameter. Pronotum ea. 2.8 times asbroad as head witheyes and 1.5 times asbroad aslong, its surface very strongly transverselyshagreened, dull. Pair of basal impressions of pronotum veryshort, 15-16 times shorterthan length of pronotum. Pronotaldisk with dense, somewhat elongate, fine punctation (average distance between punctures 52 somewhat less thantheir diameter). Posterior comers of pronotum more or less rectangularly obtuse. Scutellum transverse,with fine punctation andfinely shagr eened,weakly rounded at posterior edge, ea. 1.5 times as broad as long. Elytra oblong-oval, 1.5 times as long asbroad, their surf ace more or less shining, with very dense (on the average onlyvery slightly finerand denser thanon pronotum),transversely rugo se punctation. Prostemumbe fore procoxae ea. 2.5 times shorterthan coxal longitudinal diameter. Prostemal process very strongly developed, at first distinctly broadened posteriorly, then triangularlynarro we d toward apex; the partof prosternalpr ocess projectingbeh ind posterior edge ofprocoxae a little longer thanor equal to its maximumwidth (Plate9: 5). Impression inmiddle partof me sostemum strongly developed andoccu pying not less than2/3 extent ofmesostemum. Longitudinal suture ofmetastemum ea. 0.5 as short. Metepistemaea. 2.7- 2.8 timesas long asbroad, theirsurf ace withfine punctati.on and distinctly shagreened. Spurs of metatibiae subequal in length, 2.2-2.5 times as short as metatarsomere 1. Ab domen with dense and fine punctation and distinctly shagreened. Aedeagus as inPlate 9: 7-9. Body length: 3.7-4.4mm . Diagnosis: species differs from congeners in combining such characters as This coloration of theupper side; comparativelynot too strongly elongate antennomere 11 (less than2.5 times as long as 1 Othand 1.3-1.5 timesas long asbr oad); dullpron otum with highlya vague, dark patternaga inst a re d-brown background; antennomere 11 weakly infuscate at base. It differs from the closely related H. bomeensis Pie, 1912, inthe light terminal antennomere, denser punctation on the quite dull surface of the pronotum and the anteriorlight lateral spot reaching the lateral edge of the elytra (Plate 9: 3 ).
Holostroph us orientalis Lewis, 1895
We considerH. mul tinotatusPie, 1911, syn.nov., described fromTaiwan (Formosa) (holotyperestudied) only as a color variation of H. orientalis Lewis, 1895. The latter species iswidespr ead inJa pan,Kore a, it occursin eastern China and pos sibly inNorth Vietnam.
?Holostroph us aureofascia tus (Pie, 1954), comb. nov.
Eustrophin usaureof asciatus Pie, 1954b, Bull. Soc. Ent. Mulh.: 63 This species has been describe d from Kuat un, Fukien, China as Eustrophin us aureofasciatus(see Pie, l 954b). Judge d from some characterstaken from its original 53 description (shape of paired basal impressions of pronotum, coloration of elytra, geographicalrange) should ratherbe referredto Ho lostrophus.Unf ortunately, thetype could not be recovered.
Key to species of Holostroph us based on larvae (modified after Hayashi, 1975)
Plate 19: 1-10; Plate 20: 1-6 1. Posterior edge of abdominal tergite 9 weakly sinuate betweenurogomphi, each urogomphus short and, on inner side in front of base, without well-developed, sclerotized tubercle roundedat inner edge (Plate 20: 1,2).Abdominaltergite 9 in
lateral view stronglyconvex. 8 mm. Japan,Taiwan, China, Korea ...... H. orientalis Lewis, 1895 -. Posterioredge of abdominaltergite 9 strongly sinuate between urogomphi, each urogomphus longer, with a broadened basal part and a considerably thinner, unciformcurved apex(Plate 20: 3-6). A well-developed, sclerotizedtu bercle on inner side nearbase ofunp airedpart of urogomphi (Plate 20: 3, 5). Abdominal tergite 9 in lateral view more weaklyconvex ...... 2 2. Sclerotized tubercles on inner side near base of urogomphi more strongly approximateto each other, stronglypro jectingbeyond or atleast reaching theinner edge of urogomphi, distancebetween themnot more thantwice as great aslength of tubercle(Plate 20: 5). Innerap icalpr ocessof urogomphi proportionately shorter and, in lateral view, not less than 1.5 times as short as outer one (Plate 20: 6). 5.5-
7 .0 mm. SouthMariti me Province; Japan ...... H. diversefasciatusPie, 1921 -. Sclerotizedtubercles situatedon inner side nearbase ofurogomphi more strongly separated, not projecting behind contour of inner edge of urogomphi, distance betweenthem not less thantwice asgreat as lengthof tubercle(Plate 20: 3). Inner apical process ofurogomphi proportionately more elongate and,in lateral view,
not more than 1.3 times as shortouter as one (Plate 20: 4). 5 .5-6.0 mm. Kunashir Island;Japan ...... H. Jewisi Csiki, 1924
Tribe Eustrophini Gistel, 1856
Genera included: Eustroph us Illiger, 1802, Syn stroph us Seidlitz, 1898, and Eustrophopsis Champion, 1889. According to its author (Seidlitz, 1898), thegenus Eustrophinus Seidlitz, 1898 (type-species: My cetophagus bicolor Fabricius, 1792) differs fromEus trophopsis in the absence of a clear-cutexca vation/notch at apex of the prostemal process andin a 54 di:fferentstructure of themeso stemum. However, allthese characters which servedas basic for separation of both genera concerned prove to display a complete series of gradual transitions, without any distinct gap/hiatus. Dependent on the species, the excavationof theprosternal pr ocessand, consequen tly, the structureof the mesostemum correlated with the shape of the prostemal process can be strongly (Plate 11: 7), moderately or weakly developed (Plate 11: 13), up tonot expressed at all (Plate 11: 10). Therows of punctures on the elytra also exhibit a virtually complete transition fromwell-developed to almost indistinguishable, only withtraces when theelytra are viewed shiningthrough, or theycan entirely disap pear.Taking intoaccount all above variations, we consider thesegenera asstrict synony ms, syn.nov. The monotypic genus Curtcustroph inus Pie, 1952 is the member of the fam ily Tenebrionidae (see below).
Genus Eustroph us Illiger, 1802
Plate 10: 3-14 Type-species: My cctophagus derm estoidcs Fabricius, 1792 (Plate 10: 3-7) Thegenus includes fourspecies distributedin Europe, in the southof theRus sian FarEast, in Japan,South China and North.Amer ica. Therecent record of genus in this Turkey(W. Schawaller, personal communication), couldact uallybe attributedeither to Eustrophus dcrm cstoidcs (F.) or to an apparently new species unknown to us. Species develop in arborealfungi, e.g. Lactiporussul phurcus.
Eustroph us yunnanensis sp. nov.
Plate 9: 18 Holotype female, China, Yunnan, Jizu Shan (25.58°N, 100.21°E), 2500-2700 m, 6-10.7.1994, VitKub anleg. Thetype is kept in theN aturhi storisches Museumin Basel. Description. Oblong-oval, dark brownish with somewhat lighter elytra and abdomen; maxillarypal ps, antennae and atleast partly legs from belowred-brown, upperside coveredwith reddish-brown, more or less denseand d ecumbent pubescence. Joint 4 of maxillarypalps elongate, more or less parallel-sided,ea. 1.4 timesas long as broad andea. 1.8 times longer thana triangular joi3rd nt, latter subequal in length andwidth. Width of frons between strongly emarginate eyes ea. 2.3 times less than transverseocular diameter . Antennae when stretchedback along body not surpassing baseof elytra, antennomere 1 fairlywide and stronglyelongate, not less thantwice as long as broad; joint 2 ea. 1.5-1.6 times as long as broad and twice as short as lst; antennomere 3 elongate-triangular, ea. 2.0-2.2 timesas long asbroad and 1.8-1.9 times 55
3 1
5
12 4 <;} 15 13 10 14 6 17 2 19
Plate9. Detailsof structureofH olostrophusand Eustrophus spp. original). 1, 2 -Holostrophus bifasciatus; 3-10 -H.similis; 11-17-H.diversc fsscia tus; l8, 19- Eustroph(all usyut1DJ1L1tmais. 1, 3, 18 - pronotum and elytra; 2, 4, 14, 19 - antenna; S - prostemum; 6 - punctation of sides of prostemum (x 20); 7 - aedeagus (dorsal view); 8-apex of aedeagus (dorsalview); 9 -penis (dorsal view); 10, 13 - segments 2-4 ofmaxillary palps ; 11 - wing; 12 - metendostemite; 15 - abdominal segment 8; 16 - genital segment of abdomen; 17 - ovipositor. 56 longer than 4th; latter ea. 1.3-1.4 times as long as broad; joints 5-10 more or less distinctlybroadene d; antennomere 5 barely longer but considerablybroader than 4th, weakly longitudinal;jo int 6 subequal inlength to 5th and ea. 1.15 times asbroad as long; joints 7-10 transverse, similar in length and width, each ea. 1.2-1.3 times as broadas long ; antennomere 11 oblong-ova.4 constrictedbehind middle, with a narrowly rounded apex, ea. 1.7- 1.8 times longer than 1 Othand 1.6-1.7 times as long as broad (Plate 9: 19). Head with dense and finepunctation and finely shagreened. Pronotum rather weakly shining, ea. 3 times broader thanhead with eyes and ea. 1.85 times asbroad aslong, strongly convex, withbroadest at base whence strongly and roundlynarro wed anteriorly. Posterior comers of pronotum more or less roundly rectangular, distinctly projectingand somewhat envelopingthe base of elytra. Base of pronotum not margined at all, its sides and anterior edge fully margined. Pronotum with dense,rather medium-sizedpunctation (distancebetween punctures 0.3-1 diameter of a puncture); spaces between punctures finely shagreened. Pair of basal impressions of pronotum veryweak, considerablyshifted from posterior edge, each representedby 3-5 fairly large punctures. A fine row of punctures situated along the very posterior edgeof pronotum. Baseof pronotum shallowly bisinuate andwith a roundlypro jecting lobebef orescutellum. Scutellum weaklytransverse, roundly triangular, with moderately dense andfine punctation. Elytra shining (without distinct shagreen), oblong-oval, ea. 1.75 longer thanmaximum width,narro distinctly wed posteriorly, athumeri very slightly narrower than posterior edge of pronotum. Each elytrum. with 11 rows of punctures mainly representedby rather fine punctures elongated in longitudinaldirection (consi dera!Jly longer than on pronotum). Distance between rows of punctures not less than 3-4 transverse diameters of a puncture in row. Punctures on spaces between rows of punctures considerably smallerthan those in rows and onpronotum, moderately dense. Sides of prostemum (hypomera) withdens e and coarsepunctation, thepunctures fusing to form longitudinal wrinkles. Prostemum in front of procoxae ea. 2. 7 times shorter thanlongitudinal coxal diameter. Prostemal process margined, triangularly narrowed posteriorly, near midway ea. 3 .2 times narrower than coxal transverse diameter; process only slightlynot reachingthe posterioredge ofprocoxae. Mesosternal process slightly surpassingmidway of mesocoxae. Metastemummore or less shining, markedly coarselymore punctate laterallythan medial ly, its mediallongitudinal suture a breakbut extending posterior anteriorpart of metastemum. with small from part to meso- andmetacoxae ea. timeslonger st visible abdominal Latterbetween 1.4 than 1 stemite. Metaco:xae conspicuously separat:ed st.emite. Metatibiae su by abdominal bequal inlength to meta tarsi,their outer surfac e with13-1 4 transverse rowsof strongspines. Spurs of metatibiae distinctlybut not strongly differing in length, longest among them ea. 2.5 timesshorter than metatarsomere 1, latter somewhat shorter than followingall joints combined, joint 2 slightly longerthan 3rd and considerably broader and hardly shorterthan 4th. 57
4 6 2 G 5 D
'· ' I
17
2AC
18 8 7
Plate 10. General view and details of structure ofHallomeninae and Eustrophinae (all original). 1 -Ha llomenusbin otatus, 2 -Mycetoma 3-7 -Eustropb usdermesto ides; 8-14 nipo nicus suturalis; -E. Lewis, 1895; 15-18 - Synstrophusmacro phtha/mus. l, 11 - abdominal segment 8 of male; 2, 12 - genital segment ofmale abdomen; 3, 15 - general view; 4-labium; 5 -mandibles; 6 - maxilla; 7, 8 - antenna; 9 - wing; 10 - metatibia andmetatarsus ; 13 - lateral edge of mesostemum andmetastemum;
14 - prostemum; 16 - pregenital and genital segments of abdomen; 17 - aedeagus (dorsal view); 18 - metendostemite; a - area of anterior part of metepistemum; b - metepistemum. triangular 58
Body length: 5.5 mm. Diagnosis: species differs wellfrom congeners by the complete absence of This edgingat the base of pronotumand theslightly more strongly approximateeyes, with thedistance between them ea. 2.3 timesless than the transverse diameter(while ocular inthe other speciesthis index is less than2).
Genus Syn strophus Seidlitz, 1898
Plate 10: 15-18 Type-species: Eustroph usmacroph thalmus Reitter, 1887 (Plate '10: 15-18) The genus includes fivespecies distributed in China, Japan, the Oriental Region andNorthAmerica. Based on thestructure of theprothorax and someother characters, theonly Nearctic member of this genus, S. repandus(Hom, 1888), seems the most disjunct.Because of this,it mightprove to deservepromotion to a genus-grouptax.on ofits own.
Genus Eustrophopsis Champion, 1889
Plate 11: 1-14 'fype-species:Orch esia quindecimm.aculataLaportede Castelnau, 1840 (Plate 11: 1-4) Eustroph inusSeidlitz, 1898 (type-species:My cetophagus bicolor Fabricius, 1792), Naturg. Ins. Deutschl., 5, 2: 438, 440, syn. nov. Basedon thedescription, it seems highlyprob ablethat also themonotypic genus Pseudorchesia Fairmaire, 1883 (type-species: nigrosignata Fairmaire, 1883), P. describedfrom Misiones, Argentine is a senior subjective synonymof Eus troph opsis Champion, 1889, but unfortunately, we have failedto receivethe type for restudy. Eustrophopsisisrepresented in the world faunaby 55-56 speciesdistributed in the
NeotropicalandAfrotropical (including Madagascar) regions aswell as inthe Nearctic whereits comparati vely poor knownto occur. Itis remarkable tbatE ustroph opsis fiwna is seems to beabsent both from theOriental Region andPalearctic. Itmay be so thatit is replacedthere by speciesof thegenera Holostroph usandSynstrophusunknownfrom the Neotropical and Afrotropical regions. The Nearctic is the sole realm where representatives,however few, of allthree genera of thetribe occurtogether . Larva asin Plate 21: 1-10. 59
Eustroph op sis similis sp. nov.
Plate 11: 6-8
Holotype female, Lulua, Kapanga, Congo Beige, XI.1932, G. F. Overlaet leg. Thetype is keptin theMuseum national d'Histoire naturelle in Paris. Description. Red-brown; mesostemum, metastemum, femoraand tibiaeas well as antennomeres 5-10 andbasal part of antenna!joint 11 black; antennomeres 1-4 and apex of antennal joint 11 red-brown. Upperside covered with long, fairly dense, yellowish-greypub escence. Antennae short and surpassing posterior edge of pronotum only by their apical joint; antennomere 1 subequal in length to two followingjo ints combined; joint 2 short,ea. 1.3-1.4 times as long asbroad and ea. 1.15-1.2 times shorterthan 3rd; latter ea. 1.5-1.6 timesas long asbroad and ea. 1.3-1.4 timeslonger thana weaklylongitudinal 4th one;joi nts 5-11 distinctly broadened; antennomere 5 weakly longitudinal, ea. 1.6 timesas broad asand ea. 1.4-1.5 times longer than4th; jo ints 6-10 similarin size, each a littleshorter than 5th and 1.2-1.4 times as broadas long, joint 11 oval, ea. 1.5 times longer than 1 and 1.25-1.3 timesas long asbroad (Plate 11: 8). Eyes separatedby a Oth verynarr ow patch of frons, markedlynarr ower thanwidth of antennaljoin t 2. Pronotum convex, ea. 2. 7 timesas broad ashead with eyes and2 times asbroad as long, with rectangularly obtuse posterior comers, its base bisinuate and distinctly margined in sinuateplaces, basaledging disappearing toward sides ofpronotum; medial lobe more or less straight truncate at posterioredge (Plate 11: 6). Pronotal diskweakly shining, withfine punctation and very finely shagreened, distance between punctures in longitudinal direction ea. 1 diameter of a puncture. Pair of basal impressions of pronotummore weaklydeveloped, each one representedonly by several largepunc tures. Scutellumroundly triangular (ea. 1.2 times asbroad as long), finely and densely puncturedand distinctly transversely shagreened Elytraconvex, oblong-oval, ea. 1.45- 1.5 times longer that maximumwidth (Plate 11: 6). Each elytrum with 10 rows of punctures(regard less of a shortenedpresutural row), formedby rathermedium-sized punctures, distance between these 3-4 times as much as diameter of a puncture in rows. Rows of puncturesalmost not weakenedtowards apex of elytra. Spaces between puncturesrather weakly shining,with fine, moderately dense punctationand very finely shagreenedsimilar to thanof pronotum; size ofpunctures subequal to thaton pronotum andea. 1.5-2.0 times less thandiameter of punctures in rows. Epipleuraof elytradistinct up to lastvisible abdominal sternite. Prostemumbefore procoxae ea. 3 times shorter than coxal longitudinal diameter. Prostemalprocess stronglynotched at posterioredge, far fromreaching the hind edge ofprocoxae, its sides marginedover most of its extent (except forthe very top), more 60 or less parallel in posteriorpart (Plate 11: 7). Prosternalprocess nearits midway ea. 2,6 times narrower thancoxal transverse diameter. Metepistema ea. 2.3 timeslonger than maximumwidth. Spursof metatibiae subequal in length, 3 times as short as joint 1 of metatarsus; tarsomere 2 subequal in length to4th. Abdomen with fine punctation, distinctly and finelyshagre ened. Body length: 5.8 mmregardless of strongly declinate head, 6.9 mmtogether with head. Diagnosis: species differsfrom congeners in combiningthe following characters This first ofall: prosternal process deeply notched at posterioredge; coloration ofupp erside and that of antennae; well-developed elytral rows of punctures; very strongly approximate eyes, etc. E. similis sp. nov. is similarto Eustrophin usJesn ei Pie, 193 7 fromMozambique but, unfortunately, the typecould not be received forrevi sion. The holotype of E similis sp. nov. bears the following labels: "genre Curteustrophinus Pie, possible", " overlaeti Pie" (handwrittenby Pie himself),"L ulua, Kapanga, Congo Beige, XI.1932, G. F. Overlaet" (printed), and "type " printed on a red square.However, the status of Curteustrophin us overla etiPie, 1952 requires special attention (see below). Due to several observations, the female holotype of E. similis sp. nov. has been designatedas the type of Curteustrophin us overlaetiPie, 1952 apparently by someone else, not Pie himself. Regrettably, the original description of overlaeticontains no indication whatever, how many specimens have served Pie as the type series. So the word "possible" on one of the labels suggests that Pie could have felt inclined to exclude thisparticular specimen from the type series. Secondly, even the short diagnosis of this species as given by Pie (1952) contains some discrepancies from the above female.Thus, the holotype of E. similis sp. nov. is more uniform inelytral coloration as mentioned by Pie; it four, not three, clarifiedbasal antennomeres; its eyes are has almost contiguous, verypoorly separated, on the frons, while Pie stated that the eyes are a little approximate anteriorly; the size of the female is 5.8 mm discarding the strongly declinate head, 6.9 mmtogether with the head, while Pie gave only 7 mm mentioning nothing as to how he measured the length. Besides this, the original descriptionof overlaeti states explicitly the elytra aredevoid of rows of punctures C. that (Pie, 1952). illtimately, thetrue type of C. overlaeti been located in the collection has of theMusee Royal de l'Afrique centrale,Tervuren (see below). Inother words, there are anumb er ofvery strong indications that the type specimen of E. similis sp. nov. does not actuallybelong to the type series of Curteustrophin us overla etiPie dealt with.
Eustrophopsis sexmacula ta sp . nov.
Plate 11: 9-12 Holotypemale, Mexico,Teiu pilco, Temescaltepec, ea. 4000 ft., 1933, H. E. Hinton & R. L. Usinger Collectors. 61
11 12 CtfJ
2
8
7 10 14 13 9
Plate 11. Generalview and details of structure of J::,Ustrophopsis spp. (1-4 - afterWe grzynowicz. 1995, others original). 1-4- Eustrophopsis quindccimmacuhJta.;5 - E subaxillaris (F airmaire, 1869); 6-8 - E similis, 9-12- E se}(II13Cu.Wa;13 -14- E ochiacca. 1 - general view;2-wing ; 3 - generalview (ventral); 4, 8, 14 - antenna; 5- head (frontalview) ; 6, 9 - pronotumand elytra; 7, 10, 13 -prostemal process; 11 - tegmen; 12 - penis. 62 The type is keptin theNatural HistoryMuseum in London. Description. Elytrared-brown, each with three large black spots situated infore, middle andhind part of elytra respectively, apicalspot being more or less connected with a blackened suture; head and pronotum red-brown, latter with two black longitudinalspots bending toward medial andreaching baseof pronotum (Plate part the 11 : 9). Underside mainly red-brown, only sides and tibiae distinctly infuscate; antennomeres 1-4red-brown, joint 11 reddish-brown. Maxillarypalps infuscate with clarifiedspices ofjoints. Body oblong-oval,more or lessconvex, covered withdense decumbent yellowish-grey against a light background and black-brown or hairs: blackishagainst a darkone. Head with small densepun ctures; eyes verystrongly approximateother, toeach almost contiguous, distance between them much less thanwidth of antennomere 2. Antennae whenstretched back along body surpassingbase of elytra by ea 4 apical joints,reaching ea. anterior 1/5 lengthof elytra; antenna!joint I ea.2.4 timesas long as a shortened2nd; omere antenn 3 stronglyelongate, ea. 1.6-1.7 times as long as 2nd and its ownwidth; joi nt 4 slightly broadened and subequal in length to 2nd; joints 5-11 fairlystrongly broadened, jo int 5 ea. 1.6-1.7 timesbroader andlonger than4th, about aslong asbroad; ant ennomere6 barely shorterconsiderably but broader than 5th, both subequal length;in join ts6- 10 onlybarely shortened toward antennalapex, 1.2-1.3 5 timesas broad aslong; joint 11 oval, 1.3-1.4 timeslonger than I Oth1. and 25-1.3 times aslong asbroad. Pronotumea. I.75 timesas broad as long, its base distinctly but shallowly bisin uate, marginedin middlepart, posterior comers of pronotumrectangularly obtuse, anterior ones entirelyrounded. Medialof part pronotum sh allowly sinuateposteriorly. Pairof basalim pressionsvery weak and eachrepresented only by severalpunctures . Pronotal surfacemore or lessshining, microsculpture very weak, punctationfine and moderately dense, distance betweenpunctures up to 1-2 diameters of a puncture. Scutellum transverse (ea. 1.7- 1.8 times as broad as long), more or less straight atposterior edge, :finely puncmred. Elytra oblong-oval, a with distinctlyreflexed truncate edging,their length ea. 1.75-1.8 timesas asmaximum width; elytrashining, latmil great virtuallynot shagreened, theirrows of punctures marked, distance betweenthem ea. 1-1.7 diameters of a puncturein row. Spacesbetween rows of punctures covered with very moderately densepun s, latterbeing ea. 3-3.5 timesless thandiameter small, cture of punctures in row. Prostemum before procoxae ea. 2.5 times asshort astheir longitudinal diameter. Prostemalprocess narro w, shmply roundedat apex, not sinuate, non-marginedonly in apical 1/4 extent, ending betweenposterior 114-115 extent of procoxae. Mesocoxae distinctlyseparated. Metastemum more or lessshining, moderately finely and densely punctured, smoothin posteriorpart, not shagreened.Metastemum between meso- and metacoxae ea. 1.2 timesas long as Ist visible abdominal stemite. Metatibiaewith 11 transverserows of strong spines, their spurs being subequallength, in ea. 3.0 times 63 shorter than metatarsomere 1, latter considerably longer than two followingjoints combined. Abdomen shining, with dense and rather fine punctati.on, its stemite 1 subequal in lengthto 2nd. Aedeagus as inPlate 11: 11, 12. Body length: 6.5 mm. Diagnosis: 1bisspecies differs from congeners in the characteristic colorati.onof theup perside,the shape of theprosternal pr ocess, etc.
Eustrophopsis ochracea (Motschoulsky, 1872), comb. nov.
Plate 11: 13, 14 Eustrophusoc.hraccus Motschoulsk:y, 1872, Bull. Soc. Natural. Moscou, 45, 2: 42 Eustrophinusbo mbinus Seidlitz, 1898, Naturg. Ins.Deu tsch!., 5, 2: 441,442, syn. nov. We have studied the holotype of Eustroph us oc.hraccus Motsch. with tµe label reading "Brasilia", keptin the Motschoulsk:yCollection of the ZoologicalMuseum of the State University of Moscow, and the lectotype of Eustrophin usbom binus Seidl. (designated herewith), housed in the Naturhistorisches Museum in Vienna. Their complete synonymy beenestablished, syn.nov. 1bisdiffi was cult to reveal earlier has due to thehighly deficientdescriptions of bothspecies.
Genus Curteustrophinus Pie, 1952
Type-species: ovcrla cti Pi , 1952 C. e Basedon ourrestudy ofthe true of species (seeabove under Eustroph opsis type this similis sp. nov.), kept in the collection of the Musee Royal de l�Afrique centrale, Tervuren, the genus Curtcustrophinus appears to actuallybelong in Tenebrionidae. The originaldescription of ovcrla cti by (1952), howeverdeficient, fully C. Pie agrees with thetype at hand,including the broad body, considerablyseparated eyes, absence of elytral rows of punctures, etc.
Acknowledgments
We are most thankfulto the followingpersons who helpedus in gettingcomparative material fo r study: Dr. N. Berti andDr. S. Boucher(Museum national d'Histoire nature lle, Paris,France); Dr. M. Brendell (The Natural HistoryMuseum, London, U.K.); Dr.Y. Bousquet (Centre forLand andBiological Resources Research, Ottawa, Canada);Dr. R A. Crowson (The University of Glasgow, U.K.); Dr. R Danielsson and Dr. M. 64
Sorensson(Department of Zoology, Lund, Sweden); Dr. G.N. House (Smithsonian Institution, Washington D.C., U.S.A.); Dr. M. Janssens,Dr. E. De Coninck, Dr. M. De Meyer (Musee Royal de l' Afrique centrale, Tervmen, Belgium); Dr. J. Jelinek (NarodniPrague, Muzeum, ChechRepublic); Dr. MS. Kelley(Musemnof Comparative Zoology, husMassac etts,U.S.A); S. Kmbatov (NIICHSZR,Moscow); Dr. J. Lawrence (CSIRO,Division ofEntomo logy, Australia); Dr. l LObl (Museumd' Histoire Canberra, naturelle, Geneva, Switzerland); Dr. 0. Merkl (Natural HistoryMuseum, Budapest, Hungary); Dr. T. Nakane (Nirenokidai, Chiba-shi, Japan); Dr.A. Provonsha(Purdue University, West Lafayette, U.S.A.); Dr. W. Schawaller (Staatliches Museum fiir Naturkunde in Stuttgart, Germany); Dr. H. Schonrnann (Naturhistorisches Museum Wien,Vienna, Austria) ; Dr.H. Silfverberg(Zoo logicalMuseum, Uni versityof Helsinki, Finland); Dr. R Toyoshima (Hiroshima, Japan); Dr. W. Wittmerand Dr. M. Brancucci (Naturl:ristorischesMuseum Basel, Switzerland). We areobliged toPensoft for editing and publishing paper. Publishers this
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Kirby W. 1837. Fauna Boreali-Americana. Vo l. 4. Insects.London, J25 p. LacordaireJ.T. 1859. HistoireNaturelle des Insec�.Genera des Coleopteres. .. , 5. Paris,Librairie Encyclopediquede Roret, 750 p. Latreille P. A 1807. Genera Crustaceorum et Insectorum, secundem Ordinem Naturalemin FamiliasDisposita, Inconibus Exem.plisquePlurimlis Explicata. Vo l. 2. Paris,Amand Koenig, 280 p. Latreille P. A. 1809. Genera Crustaceorum et Insectorum, secundem Ordinem Naturalem. in Familias Disposita,Inconibus Exem.plisque Plurimus Explicata.Vo l. 4. Paris,Amand Koenig, 379 p. Lawrence J. F. & Newton A F. 1995. Families and subfamilies of Coleoptera (with selected genera, notes, references and data on family group names). Biology, Phylogenyand Classifi cations of Coleoptera. Papers Celebratingthe 80thBirthday of Roy A. Crowson. Vo l. 2. Warszawa: 779-1092. Leach W. E. 1815. Entomology. In: Brewster (ed.).Edinbmgh Encyclopedia, 9 (1): 57-172. LeConte J. L. 1866. :t-;"ew species ofColeopteraof North America. Vo l. 1, 177 p. LeConte J. L. 1874. Descriptionsof new Coleopterachiefly fromPacific slope of North America. Trans. Amer. en.tom. Soc., 5: 43-72. 66 LeConteJ. L. 1875. Descriptionof newColeoptera of theUn itedStates with notes on the geographicaldistribution. Ibid , 5: 169-208. LeConteJ. L. 1877. Descriptionsof new speciesby L. LeConte,M. D. John Proc. Amer. phil. Soc., 17: 593-627. LeConte J. L. 1880. TheCol eoptera of theAlpine Rocky Mountain. Bull. U.S. Geogr. Geol Surv., 5: 499-520. Lewis G. 1895. Onthe Cistelidae and otherHeteromerous species of Japan.Ann. Ma g. Na t Ser. 6, 15: 250-278. Hist, MelsheimerF. E. 1844.Descriptions of new species of Coleopteraof theUn ited States. Proc. Ph il.AcadNa t Sci, 2: 98-118. Miyatak.e M. 1955. Descriptionof a new species of theTetrato midae fromJapan (Coleoptera). Trans . Sh ikoku mtom. Soc., 4 (5/6): 93-94. Miyata.kc M. 1960. The genus Piscnus Casey and some notes on the family Tetratomidae(Coleoptera). Ibid , 6 (8): 121-135. MotschoulskyV. 1845a.Remarques sur des Coleopteresrusses. Bull. lacollection Soc. Na tural.Mo scou, 18 (1): 3-127. MotschoulskyV. 1845b. Observations smle Museeentomo logiquede l 'Universite imperialede Moscou.Ibid , 18 (3): 332-388. Motschoulsk:y V. 1872. Enumeration des nouvelles especes des Coleopteres rapportes de ses voyages parfeu VictorMotsc houlsky. Ibid, 45 (2): 23-55. MulsantE. 1856. Histoirenature lle de Coleopteres de Fran.et:.Paris, 201 p. Nakane T. 1955. New or little-known Coleoptera from Japan andits adj acent regions. Sci Rep. Saikyo Uni v., 2 (A): 24-42. NakaneT. 1963. Fragmenta Coleopterologica.Part 7: 27-30. Newman E. 1838. Entomologicalnot es. Entomol Ma g., 5: 372-402. NikitskyN. B. 1983. Un nouveaugenre dela famille de Melandryides (Coleoptera) de Chine. Bull.So c. mt Mulh ouse, Juillet-Septembre: 37-40. NikitskyN. B. 1985. The gencraMy cetolI11lMulsant, 1856 andOrche sia Latreille 1807 in theEastern Palearctic (Jnsecta: Coleoptera: Tetratomidae, Melandryidae). Scnckmbcrgiana biol., 65 (3/6): 265-277. Nikitsk:yN. B. 1988. Onthe status of Pachyoch th.esReitter, 1897 and Triphyllia Reitter, 1898 (Coleoptera, Cerylonidae,Te tratomidae). Folia cntom. Hung., 49: 153- 155. NikitskyN. B. 1989a. The beetle families Tetratomidae and Melandryidae of the USSR FarEast. ZoolMus. Moscow Uni v., 27: 3-87 (in Russian). Atrh. NikitskyN. B. 1989b. New taxaofbee tles from thefamilies Myc etophagidaeand Tetra.tomidae of theUSSR fauna. Zool. zhUI11Jll,68 (11): 56-64 (in Russian). Nikitsk:y N. B. 1992 (for 1991). News on the genus Triphyllia Reitter, 1898 (Coleoptera,Tetrato midae).Elytro IJ,5: 159-168. NikitskyN. B. 1992a. Fam. Tetratomidae. Keys to the Insects of the FarEastern USSR Vol. 3. Coleoptera. Part2. Nauka, St. Petersburg. P. 426-435 (in Russian). 67 Nikitsk:yN. B. 1993. The HairyFungus Beetles(Coleoptera, Myc etophagidae) of the FaunaofRussia and Adj acent Moscow StateUniv. 183 p. (inRuss ian). Lands. Press, Nomura S. and A. 1958. Anew subgenus andtwo new species of thefamily KatO Melandryidae from Japan (Coleoptera). Entomol Rev. Japan, 9 (1): 1-3. PanzerG.W . 1794. Faunae Insectorum Germanicae. Niimberg, 16 (17). Panzer G. W. 1797. Faunae Insectorum Germanicae. Niimberg, 23. Perris E. 1864.Description de quelques espCcesnouvelles de Coleoptereset notes diverses.Ann. Soc. entom.Fr. , 4 (4): 275-3 10. Pie M. 1911. H.Sauters Fonnosa -Ausbeute. Cantharidae, Lampyridae, Mordellidae. Deutscheentomol Na tionalbibl, 2: 188-189. Pie M. 1912. Coleopteresexotiques nouveaux ou peuconn us.L 'Ech.angc, 28: 93- 94 .. Pie M 1921. Notes diverses,descriptions et diagnoses. Ibid, 37: 1-4. Pie M. 1930. Nouveautes diverses.Mel. cxot -entomol, 56: 1-36. Pie M. 1934. Nouveaux Coleopteres de Chine. Entomol Nac hr., 8 (2): 84-87. Pie M. 193 7. Contnoutions a I' etudede la faunedu Mozam bique.Vo yage de M. P. Lesne (1928-1929). Mt:ID. . Estud Mus . Zool. Uni v. Coimbra, Ser. 1, No. 100: 1-9. Pie M. 1952. Genres et sous-genres nouveaux d'Heteromeres du Congo (Coleopteres). Rev. ZoolBot afr.,45: 253-255. Pie M. 1954a. Coleopteres nouveaux de Chine. BulL Soc. entom. Mulh ousc, Septembre-Octobre: 53-59. Pie M. 1954b. Coleopteresnouveaux de Chine. Ibid : 61-64. Quensel C. 1790. Dissertatio historico-naturalis, ignotas insectorum species continens. Lundae, Resp. C. P. Lundgard, 4: 1-20. ReitterE. 1887. Sullespecie del genere dicoleotteri. Na turalistaSicil., 6 (7): 84- 85. Reitter E. 1889. Beschreibungen der bekannten Tritomiden Japans mit Beriicksichtigungder neuen S ammelergeLnisseder HennGeorg� Lewis in den Jahren 1880 und 1881. "M"en. entom. Zcit, 8: 245-249. ReitterE. 1896. Abbildungenmit Beschreibungenzebn neuerColeo pterenaus der palaearctischen Fauna.Ibid , 15: 233-240. ReitterE. 1898. Neue ArtenColeopteren aus Europa angrund enzendenUnde m.. · Deutsche entom. Zcitschr., 2: 337-360. Say T. 1824. Keatings narrative of anexpedition to the sourceof St. Peters river, lake Wm nepeck, lake of the Wo ods under the command of Major Long 1823.
Philadelphia,8, Appendix, Vo l. 2: 268-378 (Coleoptera 47, ...) . SeidlitzG. 1898. Fam.Melandryidae. Naturg. Ins. Deutsch!., Berlin, 5 (2): 365- 680. Solsk:y S. 1871. Coleopteres de la Siberie orientale. HoraeSoc. entom. Ross., 7: 334-406. ViedmaMG . de 1966.Co ntnouci6n alconocimiento de las delarvas Melandryidae 68 de Europa(Col eoptera). Bos, 41 (2-3): 483-506. Viedma M. G. de 1971. Redescripci6n de la larva de Eustrophinus bicolor y consideraciones acerca de la posici6n sistematica del genero Eustrophinus [Col. Melandryidae].Azm. Soc. entom. Fr. , N. S., 7 (3): 729-733. WegrzynowiczP. 1995. Descriptionof the larvaand redescriptionof the adult of Eustroph opsis quindecimmaculata (Laporte, 1840) (Coleoptera: Melandryidae: Eustrophinae). GCIJ.us, 6 (3-4): 374-357. 69
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3
9 7 4
Plate 12. Details of larval structure of Tc tratoma spp. (all original). 1-5 - Tc tratoma (Abstrulf;z) ancora; 6, 7, 9 - T. (Tctratoma) Jimgorum; 8 - T. (Falsoxanthalia) desma.n:stii. I, 6, 8 - maxillaand labium; 2 - mandible; 3 - head; 4, 9 - antenna; 5 - epipbarynx;7 - middle leg. 70
1 A 3 2
7 4
,,.--- .....__ , 5 \ '\ I 'y \ \ \ ' \ 9 ' \ ' \ '' -...1\ ...... 8
Plate 13. of structure of Te tratoma sp). (alloriginal). 1, 2 Te tratomaetra (T toma) Details larval - ftmgorum; 3, 4 T. dCSil11lI't:Slii;5, 6 T. (Abstrulia) ai.Du;1-9 T. (A.)ancora. I, 4, 9 - abdominal- (FalsoxanthaHa) tergite2, 9; 3 - abdominal segment9 (lateral- view); 5 abdominal- tergitcs 8 & 9; 6, 7 urogompbus (lateral view); 8 middle leg. - - - 71
- .... , . ;. ·� ' . \. \ ..
5 �
9 GB 10
18
<$ 15 16
Plate 14. Generalview and details of larval structure of Pist:nus spp. (1-17 - afterHayashi, 1972, 18 original). 1-15 - Pisen usrofitarsis; 16, 17 - P. insignia; 18 - P. chujoi 1 - general view; 2 - head (dorsal view); 3 - same (ventralview) ; 4-ocelli; 5, 18 -antenna;6- epiph.aJ:ynx; 7- mand.J.o les (ventral view); 8 - maxilla(ventral view); 9 - labium; 10 - segment of prostemum(ventral view); 11 - hind leg; 12 - abdominalsegment 3 view); 13, 16 - abdominaltergi.te 9; 14, 17 - same (lateralview); 15 - spiracle of abdominal (lateralsegment 3. 72
. I I I I :\ � . "
6
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· · ·
4 1 7
Plate 15. Details of larval structureof Triphyllia kOt:IJigi (all original): 1 - head; 2 - mandibles
(ventral 3 - epipharynx; 4 - antenna; 5 - and labium (ventral 6 - middle leg; 7 - view); maxilla view); abdominal tergites 8 & 9 {dorsal view); 8 - abdominal segment 9 (lateral view); 9 - spiracle of abdominal segment 3. 73
. (J(,
n·� ' 6 10
\.-J a d� 1 5 ep 1 7 16 1 4
Plate 16. Generalview anddetails of larval structure of Penthejapana (allafter Hayashi. 1972). I - general view; 2 - head (dorsalview); 3 - same (ventral view); 4-same(lateral view); 5 - antenna; 6 - 1abrum (right part); 7 - epipilarynx; 8 - mandibles (ventral view); 9 - maxilla; 10 mala (ventral view); 11 - labium (anterior part. ventral view); 12 - hypopbarynx; 13 - hind leg; 14- - abdominal segment3 view); 15 - abdominal tergite 9 (dorsalview); 16- same(lateral view) ; 17 - spiracle of abdominal(lateral segment 3. 74
11
2 5
. /... ..·; ···... 3 j• 9 � ../ \.. .:, ..... 6
...-:- _- _
10 8 7
Plate17 . Generalview anddetails of larvalstructure of My ceto.ma suturalia1, ( 11 - afterViedma, 1966, othcn original). 1 - general view; 2-mandibles (ventral view); 3 - maxilla (ventraland labium view); 4 - epiphacynx; 5 - head; 6 - ocelli; 7 - antenna; 8 - middle leg; 9 - abdominal tergites 8 & 9 (dorsal view); 10 - same (lateralvie w); 11 - spiracleof abdominalsegment 3. 75
2 '·_/ 1
5 4
9
6
8 10
Plate 18. Detaili. of larval structure of Hallomcnus binotatus (1-8) and H. axillaris (9, 10) (all original). 1 - head; 2 - mandibles (ventralview); 3 - maxillaand labium (ventralview); 4- epipharynx; 5 - antenna; 6, 9 - abdominal 8tergites & 9 (dorsalview); 7, 10 - abdominal segments 8 & 9 (lateral view); 8 - middle leg. . . . ' ' , . ! • . ': 6 . ..
', �' 8 7
1 tJ
11
9 5
Plate 19. Generalview and detailslarval of structureof Ho lostrophusorit:Il talis after(all Hayashi, 1975). I - general view; 2 - head (dorsal view); 3 - same (ventral view); 4 -mandibles (ventralview); 5 - maxilla; 6- epipharynx; 7 - ocelli; 8 - antenna; 9 - prostemum. (ventral view); 10 - hind leg; I I - spiracle of abdominalsegment 3. 77
2 1
3
5 6
Plate 20. Details of larval structure of Ho/ostrophus spp. (1-4- after Hayashi, 1975, others original. 1, 2 - Ho /ostrophusorien talis; 3, 4 - H Jewisi; 5, 6 - H diversefasciatus. 1 - tergites 8 & 9 of abdomen; 2 - abdominalsegments 8 & 9 (lateral view); 3 - abdominaltergite 9; 4 - same (lateralview);
5 - urogomphi(donial view) ; 6 - urogomphus(lateral view) . 78
2
9
3
10 1
13 12 6 7
4 5 11 8
Plate 21. General viewand details of structure of Eustrophopsis quio.dccimmacula m larval (all after Wegrzynowicz, 1995). 1 - general view; 2 - head (dorsal view); 3 - same (ventral view); 4 - mandible (dorsal view); 5 - same (lateral view); 6 - epipharynx; 7 - anterior part of labium; 8 - hypophacy:nx; 9 - maxilla(ventral view); 10 - antenna; 11 - middle leg; 12 - abdominalsegments 9 &
10 (lateral view); 13 - spiracles ofmesostemum. 79 Index
Abstrulia 5, 6, 16, 17, 18, 19, 27, Eustrophini 4, 6, 13, 39, 40, 53 69, 70 Eustrophinus 5, 14, 40, 52, 53, 58, 60, affinis 37 63, 68 ainu 16, 17, 20, 27, 70 Eustrophopsis 4, 5, 6, 7, 8, 10, 14, 40, Alleculinae 34 53, 58, 60, 61, 63, 68, 77 almorensis 31 Eusttophus 4, 5, 6, 7, 8, 13, 39, 48, 53 anaspioides 5, 34 54, 55, 57' 58, 63 ancora 16, 17, 20, 27, 69, 70 Aphyllophorales 10, 35 Falsoxanthalia 5, 16, 17, 18, 20, 21, 27, aureofasciatus 4, 5, 52 69, 70 axillaris 39' 7 5 fungorum 16, 17, 18, 20, 27, 69, 70 fuscoguttata 3, 5, 18, 20, 21, 24, 25, 26 baudueri 16 fuscosuturalis 5, 34 bicolor 5, 53, 58, 68 fusiformis 34 bicoloripes 16, 17, 18, 20 bifasciatus 5, 48, 55 Hallomeninae 3, 5, 6, 7, 8, 9, 10, 12, binotatus 33, 35, 37, 39, 57, 75 15, 33, 37, 57 boleti 28, 29 bombinus 5, 63 Hallomenini 6 bomeensis 52 Hallomenus 3, 5, 6, 7, 8, 9, 12, 15, 33, brevicollis 34, 37 34, 35, 36, 37, 38, 39, 40, 57, 65, 75 Helops 32 chinensis, Hallomenus 3, 5, 35, 37 Holostrophini 3, 5, 7, 8, 10, 12, 13, 39 chinensis, Pseudoholostrophus 3, 5, 40, Holostrophinus 3, 5, 41, 42, 43 41, 42, 44, 49 Holostrophus 3, 4, 5, 6, 7, 8, 13, 15, 39, chujoi 28, 29, 71 40, 48, 49, 51, 52, 53, 55, 58, 76, 77 concolor 17, 18, 20 humeralis, Cryptophagus 27 Corticiaceae 16 humeralis, Hallomenus 33 crenicollis 16 Cryptophagus 27 impressicollis 3, 5, 40, 41, 42, 47 Curteustrophinus 4, 5, 54, 60, 61, 63 Incolia 5, 16, 17, 18, 19, 20 Cyanopenthe 3, 5, 6, 7, 11, 29, 30, innatus 5, 34 31, 32 insignis 27, 28, 71 cyanoptera 16, 17, 18, 20 Ischnoderma 39 dermestoides 54, 57 japana 31, 73 desmarestii 17, 18, 27, 69, 70 japonica 16, 20 discolor 3, 5, 40, 41, 42, 43 diversefasciatus 49, 53, 55, 77 klapperichi 3, 5, 34, 40, 41, 42, 45 diversicornis 5, 34 koenigi 28, 29, 72 Dryops 39
elongatus 29 Laetiporus 54 Eupisenus 29 lesnei 60 Eustrophinae 3, 5, 6, 7, 8, 9, 10, 12, lewisi 53, 77 14, 15, 39, 57, 68 longipennis 16, 17, 18, 20 80 macrophthalmus 8, 57, 58 quindecimmaculata 58, 61, 68, 78 Melandryidae 5, 6, 10, 34, 64, 65, 66, 67, 68 reitteri 3, 5, 31, 32 metallica 5, 29, 31, 32 repandus 58 minim.us 4, 5, 48, 49 reticulatus 5, 34 multinotatus 5, 52 rufitarsis 28, 29, 71 Mycetocbara 34 rufopubens 31 Mycetoma 3, 5, 6, 7, 8, 9, 12, 15, 33, 37, 39, 57, 66, 74 sakagutii 5, 16, 17, 19, 20 Mycetophagidae 6, 66, 67 Scraptiidae 34 Mycetophagus 5, 53, 54, 58 serricomis 34 sexmaculata 4, 5, 60, 61 nepalensis 3, 5, 18, 23, 24, 26 similis, Eustrophopsis 5, 59, 60, 61, 63 nigrosignata 5, 58 similis,Holostrop hus 4, 51, 55 niponicus 57 Stereaceae 16 nipponicus 3 7 subaxillaris 61 nobuchii 17, 18, 20 sulphureus 54 suturalis 37, 39, 57, 74 Notopisenus 3, 6, 8, 12, 27, 28, 29 Synstrophus 4, 5, 6, 7, 8, 13, 39, 53, 57, 58 obliquata 31 obliquatus 32 talyshensis 18 ochracea 4, 5, 61 tedaldi 16 ochraccus 5, 63 Tenebrionidae 5, 16, 34, 63 Orchesia 34, 58, 66 tesselata 16, i.7 orientalis, Hallo.menus 3, 5, 36, 37 Tetratoma 3, 5, 6, 7, 8, 11, 14, 16, 17, orientalis,Holostrophus 4, 5, 52, 53, 76, 77 18, 19, 20, 21, 23, 24, 25, 26, 27, overlaeti 6, 60, 62, 64 69, 70 Tetratomidae 3, 5, 6, 10, 14, 64, 65, pallens 5, 34 66, 67 Parahallomenus 5, 34 Tetratominae 3, 5, 6, 7, 8, 9, 10, 11, Paratetratoma 5, 16, 17, 19, 20, 21 14, 16 Penthe 3, 5, 6, 7, 9, 11, 15, 29, 30 Tetratomini 6 31, 32, 33, 73 thailandica 3, 5, 30, 31 Penthinae 3, 5, 6, 7, 8, 9, 11, 15, tokejii 34, 37 29, 31 Triphyllia 3, 6, g, 12, 15, 27, 28, 29, Penthini 6 66, 72 pictipennis 16 pim.elia 10 unicolor 13 Piseninae 3, 6, 7, 8, 9, 10, 11, 15, ussuriensis 3 7 27, 28, 29 Pisenini 6 variegatus 5, 34 Pisenus 3, 6, 7, 8, 12, 15, 27, 28, virgo 16 29, 66, 71 Pseudoholostrophus 3, 5, 6, 13, 34, 39 wittmeri 3, 5, 18, 24, 25 40, 41, 42, 43, 44, 45, 47, 49 Pseudorchesia 5, 58 Xeuxes 5, 6, 34, 37 puncticollis 34 punctulatus 35 yunnanensis 4, 5, 54, 55, 56
A new classification of the poorly-known mycetophilous and mycetoxylophilous beetle family Tetratomidae is advanced the scope in of the world fauna. Detailed descriptions and accounts are given, and keys arepresented both to adults andlarvae, so far as known. Numerous taxonomic novelties are proposed. This book is basic for a complete revision of the family, a pattern to fo llow when preparing any coleopterological treatise.
Prof. Gleb S. Medvedev Presidentof the RussianEnto mological Society