Elytra, Tokyo, 35(2): 507-527, November3,2007

Nipponocyphon,aNewGenusof JapaneseScirtidae(Coleoptera) and itsPhylogeneticSignificance

J ohn F. LAWRENCE

CSIROEntomology,GPOBox1700,Canberra,ACT2601, Australia*

and

Hiroyuki YosHIToMI

Bioindicator Co., L td. (Sapporo Branch), Kita 1, Nishi 2-11, Chuo-ku,Sapporo,060-0001Japan

Abstract Nippor1oyp11on nakane1gen et sp nov., isdescribed fromJapan, and anewsupergenericclassificationofScirtidae isproposedbasedon cladistic analysesof 22scirtidgeneraandonegenuseach fromDerodontidae,Eucinetidaeand Decliniidae with 56 adult morphological characters. Nipponocyphoninae and Stenocyphoninae, subfam nov., are proposed based on the genera Nipponoyphon and Stenocyp11on, respectively.

I nt roduct ion The family is a relatively large cosmopolitan family containing34 de- scribed genera, but the fauna is poorly known for any but theHolarctic region and phylogenetic relationships ofdescribed generaarepoorly understood. Except for the placement ofAmplectopusSHARP(seeDiscussionbelow),monophyly of the family has neverbeenquestioned,partlyduetouniquefeaturesofthemalegenitaliaandofknown larvae; however, in the recently described Stenocyphon sasaJii LAWRENCE, 2001 both male and female genitaliaexhibit a radically different structure. The discovery of an unusual scirtid-1ike from the mountains of Honshu and Shikoku, Japan, stimu- latedtheauthors tocarefullyreconsider family limitsandrelationshipsamonggeneraof Scirtidaeand relatedgroups of basal Polyphaga.

M aterials and M ethods The following abbreviations have been used for specimen repositories: ANIC -

*Mailingaddress: 130 HartwigRd., Gympie,QLD 4570, Australia

508 Joh n F. LAWRENCEand Hiroyuki YosHIToM1

f , 一 , f ,

、 ' f . . . ' -, , , - . - 一 ' f

11

Fig. 1. Nipponocyp/1on nakatlei sp nov., holotypemale. Photomicrograph digitally enhanced

A ust ralian National Collection; EUM - E him e University, Matsuyama; NMW- Natura1History Museum, Wien; NSMT - Natura1 Museum of Natureand Science, Tokyo; NZAC - N ew Zealand Collection; SEHU - Systematic Entomology, HokkaidoUniversity - T. NAKANE Collect ion. Imagein Fig. 1wasenhanced using Auto-Montagesoftware version 4.00 (Synop- tics Ltd., http:/ /www.syncroscopy.com). Symbols used in measurements and ratios: TL=total length includinghead; BL= body length(excludinghead,PL十EL);PL=pronota11engthatmidline; PW=greatest pronota1 width; EL=elytra11ength along suture, including scutellum; EW=greatest elytra1 width. The termsmesoventrite and metaventrite have been used in place of mesosternum and metasternum followingLAWRENCE(1999). Wing vein terminology follows that of KUKALovA-PEcK and LAWRENCE(1993, 2004). NewScirtid GenusNipponocypho,1 fromJapan 509

Figs 2-11. Nipponocyphot1 ,1akanei sp nov., paratype, female; 2, head in ventral view; 3, labium; 4, mandible; 5, maxilla; 6, antenna; 7, prosternum;8, scutellum; 9, metendosternite; 10, mesoventrite and metaventrite; 11, setae and punctures onelytra.

Nipponocyphon gen n o v

Type species: Nipponocyphon nakane1 sp n o v . Description. Adult. Body (Fig. 1) moderately elongate and parallel-sided, some- what flattened, clothed with suberect hairs. Head moderately strongly declined but visible fromabove;eyesmoderately large,strongly protuberant, finely facetted;sides of head immediately behindeyesslightly inflated; transverseoccipital carinaabsent; ventral epicranial ridges well developed; supra-antennal carinae well marked but not produced 510 John F. LAWRENCEand Hiroyuki YosHIToMI

over antennal insertions, which are slightly exposed; antennal fossae moderately well developed,eachextending laterally toedgeofeyebut not as wideasantennal insertion, cont inued ventrally as broad, subantenna1 groove; subocular car ina absent. F r ons barely sloping anteriorly; frontoclypea1 suture well impressed, slightly curved, with distinct tentorialpitsat eitherend; clypeusstrongly transverse,slightlywider anteriorly, sidesslightlyrounded,apextruncate. Ventralportionofhead(Fig 2) betweensubgena1 carinae flat to somewhat concave, except for gular area, which isslightly convex; gular sutures widely separated; corporotentoria1 bridge very broad. Antennae (Fig 6) relativelyshort, ifextendedposteriorly reachingjust beyondelytra1bases,moniliform to slightly incrassate, antennomeresspiculateand pubescent; 1 and 2 short and ovate, 1 slightly wider than2,3 elongate andslender, about 2.67X as long as wide, 4 slightly wider and abou t 2.OX as longas wide,5 to8progressivelyshorter and ofequal width, 9and10 about as long aswide, 11about 133X as longas wide. Labrum completely exposed, strongly transverse, about 0.37X as long as wide, sidesstrongly rounded, apexsubtruncate; tormaeshort and quadrate with acutemesal pr ocess. Mandibles (Fig 4) slightly longer than w ide at base, apex strongly and abruptly curved mesally and unidentate; basal half of outer edge with sharp dorsal carinadividingexposedsurface from that concealedbeneath clypeusand labrum;molae well-developed, asymmetrical, surfaces o f bo th ver y finely tuberculate. Prostheca well-developed, consisting of apical tuft of hairs and membranous lobe with surface hairs at molar end. Maxillae (Fig 5) with galea slightly expanded subapically and setose; 1acinia with inner edge lined with long setae and apex bearing hook-like, tridentateuncus; apicalpalpomerestronglyexpandedandsecuriform. Labium (Fig 3) withmentumtransverseand trapeziform; ligulastronglyexpandedapicallyandtruncate with internal longitudinal strut; apical palpomere attached at end of preapica1one, strongly expandedandsecuriform. Cervical sclerites well developed. Prothorax strongly transverse, about 0.5X as long aswide; sidesstrongly curved and explanate; anterior edge very weakly trisinuate, almost truncate; anterior angles rounded, not produced forward; posterior angles more or less right; lateral carinae completeand finely denticulate, without raisedmargin; baseslightly produced forming broad lobe,slightlyemarginateatmidline,withdistinctmarginobliteratedlaterally; disc veryslightly,somewhatunevenlyconvexwith narrow, slightlycurved, transversebasal groove joiningpair ofsmall, deep foveae. Prosternummoderately well developed, almost as long asshortest coxal diameter; anterior edgeslightly concave; intercoxa1process long,moderately narrow andparallel- sided,moderately archedbut notextendingbelow coxae,slightly expandedand truncate at apex andslightly overlappingmesoventrite. Procoxae transverse, subtriangular and strongly projecting, with w eak longitudinal carina ending before apex; trochantin narrowly triangular, exposed, broadly attached to endopleuron, which is slightly ex- panded apically w i th a weak anterior projection. Procoxa1cavities very broadly open internally and externally, with no traceof postcoxa1, notal projections. Scu tel lu m (Fig 8) subpentagona1, with apex somewhat rounded; anterior edge New Scirtid Genus Nippo11oyphon from Japan 511

1 m m CUA2 MP3+4 +CuA1

Fig. 12. Nipponocyp11on nakane1 sp nov., hindwing abruptly raised, straight, simple. Elytra elongate, 1.85-2.25X as long as wide, only slightly wider at base thanprothorax, parallel-sided for basal two-thirds, then gradually narrowed to conjointly rounded apices; humeri well-developed, lying just abovesharp carina extending from sides of scutellum t o elytra1 apices; epipleura n a r r o w and extending almost to apex; punctation (Fig. 11) seriate, each elytron with 10 puncture rows and scute11arystriole, each interval with additional puncture row. Mesoventrite (Fig. 10) stronglytransverse;sidesmoderately oblique; anterior edge wi t h pair of narrow, slightly cur ved and strongly elevated coxal rests; discrimen complete to base o f mesoventral process, anterior portion slightly widened forming narrow, shallow groove for reception of short keel lying above presternal process; mesoventra1 process short and acute not extending to metaventrite; mesepisternum sharplyelevatedanteriorly to form narrowprocoxa1rest;mesepimeronslightly shorter, separatedby deepgroove(pleuralsuture); mesocoxa1cavitiesconfluent, open laterally (partly closed by mesepimeron); internal meso-metathoracic joint membranous; meso- coxaeconical and projecting; mesotrochantinmoderately well-developed and exposed, elongate. Metaventrite slightly transverse, about 0.75X as long as wide, slightly convex; discrimen complete to base of metaventra1process; transverse (katepisterna1) suture well developed,extendingoneach side to about middleofmetacoxa; metanepisternum subrectangular, about 3.5 times as longas wide; metepimeron not visible. Metacoxae moderately large, only slightly oblique, extending laterally to elytra1epipleura; meta- coxa1plate completebut morewell developed mesally. Metendosternite (Fig 9) with longstalk,moderatelylongarms,well-developedventrolateralprocesses,longbutbroad anterior process and well separated anterior tendons. Hind wing (Fig. 12) about 2.25 timesas longas wide; apical fieldabout 0.25 times total wing length, with threebroad pigment patches and anarrower, longitudinal one crossingr4; radial cellsubtriangular,shorter than broad, its lumenentirelyobscured by pigment, which extends beyond the cell basally and posteriorly; cross-vein r3 short, slightlycurved,arisingseparately fromr4,which isstronglycurvedandcomplete;basal portion of RP very short and straight; radio-medial loop broad; medial spur straight, 512 Joh n F. LAWRENCE and Hiroyuki YosHIToMI almost reachingwingmargin,wherethereisaslightembayment;medial fieldwith4free veins, MP3+4十CuA1, CuA2, AA3 and AA4, the last of which extends to anal fold; MP3_+4 without basal spur or cross-vein; wedge cell small, about half as long asmedial spur,obliquelytruncateatapex,withCuA1+2arisingataboutapical thirdandsubequa1 in length toCuA1; anal notch deep; AP3+4simple. Legsmoderately longandslender; trochanter moderately long; trochanterofemora1 joint oblique but well removed from coxa; femora and tibiaesubequa1in length, the former slightlyenlargedat middle, thelatterslender andonly barelyexpanded apically; tibialsurfaceswithout longitudinal carinaeorspines; tibial spursshort,simple,subequa1; tarsusabouthalfaslongastibia,tarsomere1aboutaslongas2and3combined,slightly expanded ventrally; tarsomeres2 to4morestrongly expanded ventrally forming lobes, those on 2 and 3more or less truncate, that on 4 distinctly emarginate; tarsomere5 longest, tarsomeres1-4setosebelow; pretarsa1clawssimple; empodium not apparent. Abdomen (Fig. 13) about 2.35 timesas longaswide, flattened; ventrites1-4more or lessequal in length, 5slightly longer; all freely articulated;1aterosternitessharply delimited, narrow. Tergites I-VIII lightly sclerotized; spiracles located in pleural membrane,absentonsegment VIII; tergiteVIII inmale(Fig.14) broadly truncateand densely setose apically and broadly emarginate at base, sternite VIII (Fig. 15) with median apical incision separating two broadly rounded lobes, basally with broadly rounded plate sclerotized rim. Tergite VIII in female slightly emarginate apically, basally with paired, slightly divergent, lateral struts, sternite VIII deeply emarginate apically, truncate basally. Tergite IX in male (Fig. 16) deeply emarginate apically formingpair ofwidelyseparatedlaterotergites, but fusedatbasewith tergiteX,which is t run cate and setose at apex; sternite IX (Fig. 17) elongate asymmetrical, with obliquely truncateapex andbroadly roundedbasalplate with sclerotized rim. R ectu m with 6 cuticular rings. Aedeagus (Figs. 18-19) lying on its side when retracted; penis somewhat com- pressedlaterally, curvedsothatwhenprotractedtheapexisslightlycurveddorsallyand thebasemorestrongly curveddorsally, withanarrow projection to which thesymmet- rical parameresareattached;each paramerelongandmoderatelynarrow,expanded at middle and narrowed again apically to form narrowlyrounded lobewith fewsetae at apex; apex of penis bearingseveral spicules; endopha11us with mass of longitudinally orientedsetaeor spicules. Ovipositor (Fig 21) slightlyshorter than as last two ventritescombined and2.7 times as long (excluding styli) as greatest width, widest at middle; lightly sclerotized except for baculi. Proctiger (tergiteX) lightlysclerotized, broadly roundedanddensely setose at apex; paraprocts almost as long as gonocoxites, parallel-sided, with ventral sinuate bacula slightly converging apically; proximal gonocoxites about as long as combinedwidth,withtransverse,slightlyobliquebacula,andsidesstronglyconverging apically; distal gonocoxites each about 6 times as long as wide, parallel-sided and palpiform;styli apical,expandedand truncate,eachbearing2setae,oneateach apical angle. NewScirtid Genus Nipponocyp/1on fromJapan 5 13

16

19 Figs. 13-21. Nippo,tocyphon nakatlel sp nov., paratype, male ( l3 - 19) and female (20 - 21); 13, abdomen; 14, tergite VII I; 15, sternite VIII; 16, tergite IX - X; 17, sternite IX; 18, aedeagus in lateral view; 19, aedeagus in ventral view; 20, sternite VII; 21, ovipositor. 514 John F. LAWRENCE and Hiroyuki YosHIToMI

Glender. M asculine

Nipponocyphon nakanei sp n o v (Figs. 1-21) 1)escription. M ale. Length4.3-4.45mm. Body about 2.5timesaslongas wide, moreor lessparallel-sidedand flattened,clothed withsuberect, yellow hairs. Coloration yellowto reddish-brownor dark brown; head, prothorax, undersides, antennaeand legs somewhat lighter than elytra, which may be lighter mesally than laterally. H ead (without labrum) 0.80Xas1ong as wide, more or less flattened, slightly, irregularly concaveat middle, finely anddensely punctate; eyes prominent. Antennae moderately long, extendingposteriorlytoabout basal fourthof elytra; antennomeres3- l l distinctly longer than wide, 11more than t 5 times as longaste and more than twiceas long as wide. Pronotum about 0.5 timesas longaswide,sidesexplanateandsubsinuate, widest just in front of middle; disc somewhat uneven, with transverse groove joining pair of broad, weak impressions behind anterior edge, pair of weak, paramedia1 elevations separated by longitudinal groove just in front of middle, pair of longitudinally oval impressions just behind middle, and slender transverse groove joining pair of small, sublatera1pits just in front of posterior edge; punctation fineanddense. Elytra1about twiceas longaswideand5 times as longaspronotum, with rowswith punctures which are coarser and more densely packed than on pronotum, but with rows of smaller punctures in intervals; interspaces smooth and shiny. Ventrite V moderately evenly rounded except at apex, where thereis anarrow, shallow emargination. F e m a1e. Length 4.6 mm. Antennae somewhat shor ter than in male, with antennomere111essthan t 5 timesas longasteand lessthan t 5 timesas longas wide. Body2.34timesaslongaswide. Pronotum0.51 timesas longas wide. Elytra196times as long as wide and 5.l l times as long as pronotum. Ventrite 5 evenly, narrowly rounded at apex. Measuremen ts and ra tios. Measurements in mm: TL (male) 4.3-4.45 (4.36), PL 0.6-0.68(0.65),PW1.16-1.36(1.26), EL3.12-3.6(3.4), EW144-1.88 (1.66±0.l9). Ratios: BL/EW2.21-2.63 (2.46);PL/PW0.50-0.53 (0.52), EL/EW 185-2.24 (2.06); EL/PL 4.88-5.67 (5.24). Ratioof antennomere lengths: male:1.12: 1.00: 1.75 :1.75: 1.62: 1.50:1.50: 1.50: 1.50: 1.50: 2.40; female: 1.43: 1.00:1.71: 1.57: 1.43 :1.43: 1.43 : 1.29 : 1.29 : 1.29 : 1.86. Antennomere length-width ratios: (male) 1.12:1.33: 1.75 : 1.55 : 1.44 : 1.33 : 1.33 : 1.33 : 1.33 : 1.33 : 2.22; ( female) 1.25 : 1.16 : 2.00 : 1.57 : 1.25 : 1.25 : 1.11 : 1.00 : 1.00 : 1.00 : 1.44. Type material. Holotype: Male, Nikko, Tochigi Pref., 11~13-VI-l967, H. TAKIzAwA leg. (NSMT). Paratypes:1 female, Tashiroyama-rindo, 1,300m, Kuriyama, Tochigi Prof.,6-VI-l982, N. MoRIsHIMAleg(ANIC, drywingslideand dissection in glycerine); 1male, Mt. Gomadan, Wakayama Prof., 22-V-1997, 1. MAToBA leg. (NSMT); lmale, Chu-zenji, Tochigi Pref., 9-VII-1917, Edme GALLoIs (SEHU); 1 male, Joju, lye, 1~3-VI-1967, T. KosAKA (SEHU); 1 female, Shobugahama, Nikko- New Scirtid Genus Nippo11oyphon from Japan 515 shi,TochigiPref.,11-VI-2005,S. MAEHARAleg. (NMW);1male, Mt.Jirogyu(F.I.T), Kisawa-son, Tokushima Prof., 4~13-VI-2004, K. TANAKA leg. (NZAC); 1 female, Hikawa-path, Daibosatsu Mts., Yamanashi Pref., 14-VI-1992,S. TsuYUKI leg. (EUM); 1 male,0kuyarito(FIT), Kisawa-son,TokushimaPref.,31-V~4-VI-2004, K.TANAKA leg. (EUM); 1male, Mt. Maruzasayama, near Mt. Tsurugi, TokushimaPref., 11-VI- 1972, M. YosHIDA leg. (EUM); 1 female, 0odaigahara, Kamikitayama-mura, Nara Pref., 21 ~24-VI -2005, T. KIsHIMoTo leg. (NSMT); 1male& 1 female, 0tomedani (alt. 1,260m), Higashiiya, Miyoshi, Tokushima Pref., 28-V ~5-VI-2006 (FIT), K. TANAKA leg. (EUM); 1male, Minokoshi (alt. 1,360m), Higashiiya, Miyoshi, TokushimaProf., 28-V ~5-VI-2006 (FIT), K. TANAKA leg. (NMW). Dist ribution . Japan (Honshu, Shikoku). Etymology. Thespecies isnamedafter the lateDr. Takehiko NAKANE, who gave us theopportunity to study thisspecies.

Cladistic Analysis In order to assess the phylogenetic relationships of Nippono〔:yphon to selected gener a o f Scir tidae and members of related families of basal Polyphaga, 56 adult morphological characters were coded for Nippono yphon, 21genera of Scirtidae, and one taxon each f rom the families Derodontidae, and Decliniidae (see Appendices 1 and 2). The data were initially coded in Delta format using the Delta Editor (DALLwITz et al., 2000a, 2000b) and converted to Hennig86 (FARRIs, 1988) files for use with Winclada (NIXON, 1999) and N ona (GoLoBoFF, 1999). A ll characters were treated as unordered. With all t axa and characters included and Derodontus as outgroup, a heuristicsearch using multiple TBR十TBR with95 replica- tions produced three shortest trees with a length of 169, consistency index of 41 and retention index of59. Astrictconsensus of thesetrees isshown in Fig 22 w i th B rem er support valuesgiven for themajor nodes; thesametreeshowingunambiguouscharacter changes isshown in Fig 23. Inall trees, Nycteus isbasal to threeunresolved clades:Declinia, Nippono〔1yphori and remaining Scirtidae; Stone〔typhon is basal to the remaining20 scirtid genera (hereafter referred t o as Scirtidae MP), a clade is formed with Hydrocyphon, Amplectopus, Sarabandus,SacodesandE1odes, and the followinggenericpairs are recognized:Sacodes 十E1odes,Atopida十Byrrhopsis,Macrodasc111us十PrionocyphonandScirtes十Ora. Inone of thetrees Nipponocyphon and_Declinia aresister taxa, while in theother two Declinia is basal toNippono〔:yphon or viceversa. TheDeclinia十Nipponoyphon十Scirtidae clade hasa Bremer valueof 4and issupportedbyseven synapomorphies, but only three, 39- 1 (metendosternitewith ventrolateralprocesses), 41-1 (AA4meetinganal fold) and51- 1 (loss of spiracles onsegment VIII) areuniqueand unreversed. Of the three changes supportingStenoyphon十Scirtidae MP, 52-2 (fusion of tergites IX and X) is unique and unreversed and 53-0 (absence of rectal rings) is a reversal to the condition in Derodontus. Thereappears to be more substantial support for the Scirtidae MP clade,

516 John F. LAWRENCE and Hiroyuki YosHIToMl

0 0 0 0

t、 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

、0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

l.n 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 l「、1

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 l「、1 ll'、1 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

e、0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 A s I s

、 一 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 u o 00 0 0 0 0 0

o 、 0 0 0 0 0 0 、 0 0 一 一 t 0 0 0 0 0 e' l o - P - s 、0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

」 0 0 0 0 0 JxI x oJ

」 一差 0 0 0 0 0 C'、1 '、1 ll、l 0 C、l 0 ' 、1 ( '、1 C'、l C、l ll'、1 C、l

tl'、 0 0 0 0 ' 、1 l「、l l、・1 '、1 e、l ('、1 l'、l (、l C、l e、、l (、1 e 、l . 9 ・ s - e 0 0 0 0 0 0 0 0 0 0 0 0 0 」 .I e o 9 = 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 o 」

o 一 一 一

[- n[ 0 ・〇・ 0 0 0 ) 0 、 e 0 0 0 0 0 0 0 ll'、1 0 0 0 0 0 0 0 0 e

rL er q I o - 00 e 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 l o t、 0 0 0 0 0 0 0 C'、l 0 ('、1 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

,, r、 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

l - _ 'S:. ・ :1 'S ミ. 0 517 l 0 0 0 、、 0 0 0 0 0 0 0 0 0 0 0 :: 0 0 0 0 0 0 0 0 0 0 コ l C l 、、 0 、 0 0 0 0 0 e 0 0 0 l l l 0 0 0 0 、 0 0 0 0 0 つ ' 1: C 0 0 0 0 0 0 0 0 0 0 0 l 0 、 0 0 0 0 0 0 0 0 0 l :1 ・ l 1 l 、 、 0 0 0 0 0 0 0 0 0 0 0 l e 1 e' 0 Japan 、 、 0 0 0 0 0 0 0 0 0 0 0 0 C' 'S l e 、 0 0 0 0 0 0 0 0 0 0 S o: ・ from 0 0 c c 0 c c 0 c l ll . 0 0 0 、 0 0 0 0 0 0 ・二 0 0 0 0 0 0 0 0 0 ミ ' ・要 l ll 、 0 0 0 0 0 0 0 0 0 0 l l C 、 0 0 、・ 0 0 0 0 0 0 l ( : 0 0 「、 c 0 0 0 「、 e c c l 二 l= ミ C : 1 l Nipponocyphon ' 0 0 、 0 0 0 0 0 0 0 ( . ミ l 0 0 0 0 0 0 0 、 e l 0 、 0 0 0 0 C 0 0 0 0 0 0 e e 0 Genus l 1 、 0 0 0 0 0 0 0 、、 0 0 ( e' l 1

、 0 0 0 0 0 「、 0 0 0 l 0 0 l' 0 0 0 0 0 0 0 :. 0 0 0 0 Scirtid 0 0 0 ・二 0 0 0 0 0 0 0 0 0 0 0 0 0 0 New 0 0 0 0 0 0 ミ : l 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 、、 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 c 0 0 c 0 c c 0 0 l 0 0 0 c 0 c 0 l 二 S ・ 、 、 、 、 0 e t 0 0 e ltln t 00 0 l - 、 :!:

一 一 〇・ q - o- q P o n u u o

518 John F. LAWRENCE and Hiroyuki YosHIToMl

Fig 22. Strict consensus of three shortest trees produced by NONA f rom the total data set, using multiple TBR 十 TBR (mutt*max*) with 95 replications and one starting tree per replication, showingBremer support valuesof2ormore. Length=169 (CI=41, RI=59). but itsBremersupport isonlyone. Of theeight unambiguouschanges, 15,1(lossof 1acinia1uncus), 41-1 (reduction of veins in medial field), 52-0 (parallel or diverging basalstrutsonsterniteVIII),53-1 (baseofsegment IXopen) and56-1 (fusion or loss of parameres) are unique and unreversed. Within the clade containing most scirtid genera, theonlycladeswithrelativelystrongBremersupport areSacodes十E1odes(5), Scir tes十Ora (5) and Macrodasci11us十Prionocyphon (3). The low support value for ScirtidaeMPappearstobeassociatedwiththeinclusionofAmplectopus,agenuswhich is discussed further below. When theanalysis isrun with this genus excluded and with five uninformativecharactersdeactivated, the resultingcopnsensusof 9shortest trees hasasimilar topology,butScirtidaeMPhasaBremer valueof4andacladeformed by Sarabandus十E1odes十Sacodes has a valueof 2. A final analysisexcludingDerodontus, withNycteusasoutgroup,producedthesingleshortest treeshown inFig 24; hereagain the Bremer value was4 for Scirtidae MP, with I)eclinia basal to Nippono〔:yphon十 Scirtidae, andNipponocyphon basal toStenoyphon十ScirtidaeMP. New Scirtid Genus Nippo11ocyp/1ot1 fromJapan 519

Fig 23. Cladogram in Fig 22 showing unambiguouscharacter state changes 520 John F. LAWRENCE and Hiroyuki YosHIToMl

D is cussion Phylogenetic relationships o f the family Sci rtidae and ot her members o f t he superfamily Scirtoidea ( = Eucinetoideaof CRowsoN, 1960) havebeen reexamined in several recent works. LAWRENCEet a1. (1995) includedScirtoideaat the base of the series (Dasci11iformia of CRowsoN, 1950, 1955), but since then it has become apparent through both morphological (LAWRENCE, 2001) and molecular (CATERINo et a1., 2002) studies that both Scirtoideaand Derodontidae may occupy a morebasal position within Polyphaga. In their study of the pterothorax of selected Scirtidae (species of E1odes, Cyphon, Pseudomicrocara and Ora), FRIEDRIcH an d BEuTEL(2006) failed to findsupport for abasalpositionofScirtoideawithinPolyphaga except for theabsenceof ahind wing “bending zone” inall other elateriform lineages; however a n inclusion of this superfamily within Elateriformia was only weakly sup - ported by an elongate metanepisternum, triangular radial cell and reduction of RP branchesintheapical field of thehindwing. Theabsenceofabendingzonecannot be consideredasynapomorphy for ElateriformiaexcludingScirtoidea, since this occurs at least in some Artematopodidae. However, those features used to unitescirtoids with Elateriformiaareeither unclearor just incorrect: ashort,broadmetanepisternum occurs both in the scirtoid family Eucinetidae and in some Rhipiceridae among the basal Elateriformia, theboundarybetween “rounded” and “triangular” radial cells appearsto break down in various Elateriformia, as well as in Bostrichiformia - Cucujiformia, and RPbranchesintheapical fieldarequiteobviousat least inDasci11idae. Thepositionof theScirtoidea continues to remainambiguous, and will hopefully be clarified by theuse of a larger morphological character set combined with DNA data. Thediscoveryof thefamilyDecliniidae(NIKITsKYeta1.,1994) reinforcedthe link between Scirtidae and Eucinetidae, since Declinia shared a type of aedeagus and the presenceof rectal ringswithEucinetidae,whilethewingvenationandlossof theeighth spiracles weremore reminiscent of Scirtidae. Theunusual ChileangenusStenocyphon (LAWRENCE,2001) hasa largeexposedprotrochantin likethat inDecliniaandtrilobate aedeagus with a distinct phal1obase; however the redu ced prosternum, projecting procoxae, reduced eighth spiraclesandwing venation aremoretypical of the family Scirt idae, in which the genus was placed. In Nipponocyphon, we have yet another transitional form, which, likeStone〔:yphon, lacks thetwo tibial carinaepresent inalmost all scirtidsand hasan atypical aedeagus, which lacksaphal1obasebut hasarticulated parameres. Furthermo1-e, it has the rectal rings found in Eucinetidaeand Decliniidae and 10 distinctelytra1puncturerowsplusascute11arystricto, which occur inno other sci rtid. Basedontheaboveanalyses,Nipponoyphon either liesoutsidethefamilyScirtidae or forms the most basal scirtid clade. Although it forms a monophyletic group with Declinia inoneof thethreetreesproducedwiththetotal dataset, itneverclusterswith Stone〔:yphon. The latter genus is weakly attached to astrongly supported main scirtid cladein all trees. Thepresent dataset does notsupport anystrongly supported clades New Scirtid Genus Nippono yp/1on fromJapan 52 1

Fig 24. Single shortest tree produced by Nona from the data set excluding Derodo11tus and with Nycteus as outgroup, using the same search strategy and showing Bremer support values of 2 or more. Length=153 (CI=41, RI=58).

within Scirtidaeproper, except for thegeneric pairsE1odes十Sacodes,Scirtes十Ora and possibly Macrodasci11us十Priono yphon. Currently availabledataon phylogenetic relationships within the familyScirtidae, assummarized by YosHIToMI (2005), arebased primarily on Palaearctic genera and relytoagreatextent on charactersof thelarvae. Unfortunately, larvaeareunknown for critical taxadiscussedaboveandnoneof theSouthern Hemisphere larvaedescribedby HANNAPEL and PAULUS(1991) have been associated with adults. The cladogram producedby HANNAPELandPAULUS(1987), basedon larval characters only, and that of YosHIToMI (2005), based on both larvae and adults, agree in theseparation of an E1odes十Sacodes clade from a clade containing the remaining Palaearctic genera with Hydr・o〔:yphon at its base. Thecladograms produced here contain thesame two clades, but with Hydro〔typhon attached to the former rather than the latter. Themost problematic genus within themain body ofScirtidaeis the New Zealand Amplectopus, which was moved to the family Che1onariidaeby KAsAPand CRowsoN (1975), based mainly on the fusionof the first three ventrites, and returned toScirtidae by LAWRENCEet a1. (1995). Although the reducedprosternum, typeof metendoster- niteandwingvenation inAmplectopusaretypical ofScirtidae,severaldivergent features aresimilar to thosein Declinla. These include the5-segmentedantennal club, distinctive 522 John F. LAwItENcEand Hi royuki YosHIToMl pedicelshape,dorsalmandibularcarina,apicallywidenedepipleura,mid leg impressions on metaventrite, mesepisternum and epipleura, single tibial carina, and fusion of the basal threeventrites. AspointedoutbyLAwRENcEetal. (1995),someofthesefeatures differ in detail. TheAmplectopusantenna, for instance, lacks thespecialized sensilla found inDeclinia. Furthermore, theaedeagus isofadistinctivetypefoundonly in the familyScirtidae(NYHoLM,1972,2000; YosHIToMI, 2005). Thetegmen isdorsoven- trally flattenedanddeeplyemarginate,withabroadbasebutno indicationofaseparate pha11obase, and thepenis isalso flattened, with styliform parameroids and aprostheme bearingapair ofshort lateral hooks. The information presented above presents us with the following nomenclatura1 options: 1) to in clude Declinia in a more broadly defined Scirtidae, also including Nipponocyphon and Stenoyphon, 2) to place Nipponocyphon in a new family, 3) to propose new scirtid subfamilies for both Nippono〔:yphon and Stenoyphon, or 4) to includeNippono〔:yphon inScirtidaebutmakenochangesat thesupergenericleve1. We hesitate tomake family level changes at thistime, especially when immaturestages are known for none o f these taxa. On the other hand, we feel that some changes in classification should reflect the basal positions of bothNipponocyphon and Stenocyphon and thenumber of important adult features uniting the remaining scirtid genera. We thereforeproposethe followingnew taxa: Nipponocyphoninaesubfam n o v and Steno- cyphoninaesubfam nov., for thegeneraNipponocyphon andStone〔:yphon, respectively. At thepresent time, therearenoother subfamilies or tribeswithinScirtidae, except for Atopidini proposed without justification by PIc ( l914) for the New Zealand genus Atoplda. Thesubfamilies ofScirtidaemay beseparated by the following key:

Prosternum in front ofcoxaeat least half as longaspresternal process;elytra with 10 distinct puncture rows and scute11ary striole; mandible with well developed motaandmembranousprostheca; apical maxillary palpomerestronglyexpanded apically, subtriangular; sides of pronotum broadly explanate with denticulate edges; hind wing with wedge cell; tergite X at least partly free from tergite IX; aedeaguswithphal1obaseabsent andparameresarticulated tobaseof penis. ・・・・ Nipponocyphoninae - Prosternum in front of coxae less than half as long as presternal process; elytra punctation not seriate; mandibular mota, if present, not accompanied by mem- branous prostheca; apical maxillary palpomere not or only slightly expanded apically; sides of pronotum, if explanate, not denticulate; hind wing without wedge cell; tergite X completely fused to tergite IX; parameres articulated to

・・・・・・pha11obase, fixed o r absent . ・ 2 2. Frontoclypea1suture distinctly impressed;mandiblebidentate;1aciniawithuncus; protrochantin large, quadrateandbroadly exposed; hindwingwith4 free veins in medial field; outer edge of tibia without longitudinal carina; aedeagus latera1ly compressed with distinct phaliobaseand articulated parameres Stenocyphoninae New Scirtid Genus Nipponocyp/1on from Japan 523

Frontoclypea1suture vaguely impressed or absent; mandible unidentate; lac inia wjthout uncus; protrochantin slender and usually more or less concealed by stronglydeclinedhead;hindwingwith3or fewer freeveinsinmedial field;outer edgesof tibiaealmostalwayswithpairedlongitudinalcarinae(rarelywithsingle carina); aedeagus dorsoventrally flattened, without pha11obase and with para- meres fixed or absen t Scir tinae

Acknowledgments Wewish toexpressoursinceregratitudetoDr.MasahiroSAKAI(EUM), thelate D r. Takehik o NAKANE, Dr. Masahiro OHARA (SEHU), Mr. Isao MATOBA (Wakayama), Mr. KouichiSAT0and Mr.Satoshi MAEHARA(Tochigi), Mr. Masataka YosHIDAand Mr. Koji TANAKA(Tokushima), Dr. ToshioKISHIMOTo(Tokyo),and Dr. RichardA. B. LEscHEN(NZAC) for their offeringthepreciousmaterials. CSIRODivisionofentomologyandTheAustralianNational Insect Collectionare acknowledged for allowingJFL tostudyspecimens in their care. Special thanksare giventoDr. AdamSLIpINsKI for hisassistanceinpreparingFig.1.

Appendix 1. Taxa for Cladistic Analyses AmplectopusSHARP, 1886.BasedonAmplectopusovalisSHARP(NewZealand). AtopidaWHITE,1846.BasedonAtoplda lawsoni BRoUNandAtopldasp. (New Zealand). ByrrhopslsCHAMPION,1913. BasedonB gravidus (SHARP) (NewZealand). CyphanusSHARP,1878.BasedonCyp/1anussp. (NewZealand). CyphonPAYKULL,1799.BasedonCbrevico11isLECoNTE(westernNorthAmerica),C. co11aris (GU直RIN-MtNEvILLE) (eastern North America), C concinnus LECONTE(western North America) andCyphonspp anddescriptionsandillustrations in YosHITOMI (2005). CyphotehisSHARP, 1878. Basedon C angllstifronsSHARP1878,58 (NewZealand). De(、1iniaNIKITsKYet a1., 1994.Basedon femalesofD relicta NIKITSKYet a1. (eastern Russia) andD. versicolor SAKAI etSAT0 (Japan) anddescriptions and illustrations inSAKAI &SAT0 (1996). Derodontus LECoNTE, 1861. Based onDerodontus spp. (North America). E1odes LATREILLE,1796.BasedonE apicalis LECoNTE(North America) anddescriptionsand illustrationsin YosHIToM1 (2005). Heterocyphon ARMSTRONG,1953.BasedonH australis (ERICHSON) (Australia). Hydroyphor1REDTENBAcHER,1858.BasedonH.sato1YOSHITOMI (Japan)anddescriptionsand illustrationsin YosHIToMI (2005). MacrocyphonPIc,1918.Basedon M spencei ARMSTRONG(Australia). Macrodascilhis CARTER, 1935. Based on M dentlcornls CARTER(Australia). Macrohe1odes BLACKBURN, 1892. Based on M crassus BLACKBURN (Australia). Microcara THOMSON,1859.BasedonM explanata (LECoNTE) (northern North America) and M testacea (LINNAEUS) (Europe). Nippono yphon gen nov. Based on N nakane1 sp nov. (Japan). Nycteus LATREILLE,1829.BasedOnN 可umatus(LECONTE) (North Ame「iCa). 524 John F. LAWRENCE and Hiroyuki YosHIToMl

OraCLARK, 1865.Based onOra spp. (FloridaandBrazil) anddescriptions and illustrations in YOSHITOMI (2005). PrionocyphonREDTENBAcHER,1858.BasedonP discoldeus(SAY) (eastern North America) and P rliger KITCHINGet ALLsoPP(Australia) anddescriptionsand illustrationsin YosHIToMl (2005). Pseudomi(1'oca''a ARMSTRONG, 1953. based on P. orlentalis ARMSTRONG and P. va,・1abi lis ARMSTRONG (Aust ralia). Sacodes LECONTE, 1853. Based on S. pulc11e11a (GUERIN-M色NEvlLLE) and S. thoracica (GU直RIN-MENEvILLE) (North America) anddescriptionsand illustrations in YosHIToMI (2005). Sarabaltdus LEECH,1955.BasedonS robustits(LECoNTE) (eastern North America). Sclrtes ILLIGER,1807.BasedonSclrtesspp. (North Americaand Australia) and descriptions and illustrations in YosHIToMI (2005). Stenocyphor1LAWRENCE,2001. BasedonS. sasaJli LAWRENCE(Chile). Veronatus SHARP, 1878. Based on V t1'1costetlils (WHITE) and Veronatussp. (New Zealand).

Appendix 2. Characters and Character States for Cladistic Analyses 1. Frontoclypea1suture:0,vaguely impressedor absent; 1, distinctly impressed. 2. Subocular carina: 0, absent; 1, present. This refers to a sharp ridge lying between t he subgena1 ridgeand theeye, thus formingoneedge of the subantenna1groove. 3. Subgenal ridge: 0, absent; 1, less sharply defined and located immediately beneath and behindeye;2,moresharply definedandextendingwellbehindeye. 4. Antennomere1: 0, not or only slightly inflated, not carinate; 1, strongly inflated and more or less ca rinate. 5. Antennomere2:0, not wider atbasethanat apex;1, distinctly wider at base thanat apex. 6. Antennomere3: 0, not distinctly shorter than2; 1, distinctly shorter than2. 7. Antennomere4:0,shorterthan2and3combined;1,between1and2timesaslongas2and 3 combined; 2, more than2 times as longas2and3 combined. 8. Antennomeres4 to le: 0, neither serrate nor pectinate; 1, serrate or pectinate. 9. Apex of labrum: 0, subtruncate to slightly convex; 1, slightly concave or emarginate; 2, deeply emarginate or bilobed. 10. Mandible: 0, bidentate; 1, unidentate. 11. Dorsal surfaceof mandible:0, withoutcarina fittingover sides of labrum;1, with carina fitting over sides of labrum. 12. Mesal edge of mandible: 0, without teeth or retinacula; 1, with two or more teeth or retinacula. 13. Mesal edgeof mandible:0, with membranous prostheca (sometimesaccompaniedby fringe of hairs); 1, with fringe of hairs only; 2, with neither prostheca nor fringe of hairs. 14. Mandibular mota: 0, well developed (occupying basal fifth or more); 1, very small (occupyingbasal tenth); 2, absent. 15. Lacinial apex: 0, with uncus; 1, without uncus. This refers to a sclerotized, hook-like process, usually tridentate. 16. Apical maxillary palpomere: 0, cylindrical to fusiform, not apically expanded; 1, apically expanded and subtriangular. New Scirtid Genus Nipponocyphon fromJapan 525

17. Preapical iabia1palpomere:0, not distinctlyenlargedor obliqueat apex, apical palpomere arisingfromaboutmiddleofapicaledge;1, distinctlyenlargedandobliqueatapex,apical palpomere arising near inner portion of apical edge; 2, highly distorted, so that apical palpomerearises at middle or near baseand palpi appears bifurcate. 18. Prothorax: 0, not widest anteriorly; 1, widest anteriorly. 19. Sidesof prothoracic disc:0, not orslightlyexplanate;1, distinctlyexplanate. 20. Baseof prothorax:0, notorslightlynarrower thanelytra1bases;1, distinctly narrower than elytra1bases. 21. Anterior edgeof pronotum:0, truncateor emarginate, not forming continuouscurvewith lateraledges;1,strongly rounded, formingcontinuouscurve with lateraledges. 22. Anterior anglesofpronotum:0,absentor notproducedforward;1, producedand rounded or broadly angulate;2, produced andacute. 23. Lateral pronota1carinae:0, simpleor minutelycrenulate;1, denticulate. 24. P osterior angles of pronotum: 0, absent or broadly rounded; 1, obtuse or right; 2, moderately to strongly acute. 25. Pronota1disc just in front of posterioredge:0, withoutpair ofsmall pits;1, withapair of small pits. 26. Presternal process:0, not abruptly bent, broadenedand attenedat apex;1, abruptlybent, broadened and attened at apex. 27. Presternalprocessventrally:0,extendingalmost tocoxal apex; l,endingwell beforecoxal apex. 28. Protrochantin:0,largeandsubquadrate,lyingbetweencoxaandedgeofnotum, forming part of lateral thoracicwall; 1,smallandnarrow, lyingin front ofcoxaandnot forming part of thoracic wall. 29. Anterior edgeofscutellum:0, notor graduallyelevate;1, abruptlyelevated formingsharp ridge. 30. Elytra1punctation:0, distinctlyseriate;1, not distinctlyseriate. 31. Elytra1epipleuron: 0, narrowedapically; 1, slightly widenedat apex. 32. Mesoventrite:0,dividedbylongitudinal grooveordiscrimen;1, notdividedby longitudinal grooveor discrimen. 33. Mesoventralcavity:0,present; 1,absent. This referstoadistinct depressionlyingbetween andin front of themesocoxa1cavitiesandnot justaslight wideningof themesothoracic discrimen. 34. Mesocoxa1cavities:0,contiguous;1, narrowlyseparated;2,moderatelytowidelyseparated (more than 0.4X shortest diameter of coxal cavity). 35. Apex ofmesoventra1process:0, notcleftor emarginate;1, cleft or emarginate. 36. Metathoracic discrimen:0,completetobaseof intercoxa1process;1, incompletebutmore than half median length of ventrite (excluding intercoxa1process); 2, less than half medianlengthof ventrite(excludingintercoxa1process). 37. Metaventrite,metepisternumandanterior portionofepipleuron:0,without impressionsfor housingmid legs;1, with impressionsfor housingmid legs. 38. Metacoxa1plate:0,extending to lateraledgeofcoxa;1,extendingbeyondmiddleof coxa but not to lateral edge; 2, not extending tomiddle of coxa. 39. Metendosternite:0, without ventrolateralprocesses;1, withventrolateral processes. 40. Radial cell of hindwing:0, formingequilateral triangle;1, formingelongatetriangle. 41. Medial fieldof hindwing:0, with4ormore terminalveins;1, with3or fewer terminal veins. 526 John F. LAWRENCEand Hiroyuki YosHIToMI

42. Cross-veinjoining MPl+2and MP314:0, present; 1,absent. 43. Wedge cell of hind wing:0, present; 1, absent. 44. Wingvein AA4:0, notmeeting toanal fold;1,meetinganal fold. 45. Wingvein AP3+4:0, forked toformAP3and AP4;1,simpleor absent. 46. Mesotibia:0,without longitudinal carina;1, withsinglelongitudinal carina;2, withpaired longitudinal carinae. 47. Metafemur: 0, not much wider than mesofemur; 1, much wider than mesofemur. 48. Metatibial spurs:0,moreor lessequal in length;1, greatly differingin length. 49. Number of basal ventrites connate: 0, none or two; 1, t hree; 2, four. 50. Anterior edgeofventrite1 (sterniteIII): 0, withintercoxa1process;1,without intercoxa1 process. 51. Spiracles onsegment VIII:0, present; 1,absent. 52. BaseofsterniteVIII inmale:0,withparallelordiverginglateralstruts;1,withlateralstruts meeting to formclosedbasal rim. 53. Base of segment IX inmale: 0, closed forminggenital ring; 1, open with separate basal st r u ts. 54. Proctiger (tergite X) inmale:0,completely freefromtergiteIX;1, partly fused to tergite IX; 2. completely fused to tergite IX. 55. Rectal rings: 0, absent; 1, present. These structures are illustrated in L AWRENCE et a1. (1995, fig. 17). 56. Parameres: 0, basally articulated; 1, fixedor absent.

要 約 JohnF.LAWRENCE・吉富専之l :新属Nipponocyphonの「ヨ本からの発見とその系統的位置. - 日本からたいへん特徴的なマルハナノミ科の新属新種ナガマルハナノミ Nipponocyphonnakanei gen etsp nov. を記載した. 本属を含め, マルハナノミ科22属と外群3科(Derodontidae,Euci- netidae, Decliniidae)に対して成虫の外部形態56形質を用い系統解析を行った. その結果, ナガ マルハナノミ属は南米のStenocyphon属を含めたマルハナノミ科と市妹群関係になることが判明ll し た. そこで, ナガマルハナノミ属とStenocyphon属それぞれにNipponocyphoninaeとStenocy- phoninaeの新亜科を創設した. また, 残りのマルハナノミ科に対して, Scirtinae亜科を提唱した.

References

CATERIN0, M. S., V. S. SHULL, P. M. HAMMOND& A. P. VOGLER,2002. Basal relationshipsOf C01eOpte「a inferred from18S rDNAsequences. Zoo1.Scripta,31:41-49. CRowsoN,R. A., 1950. Theclassificationof thefamiliesofBritishColeoptera (part).Ent monthlyMag., 86: 327-344. - 1955. The Natural Classification of the Families of Coleoptera. 187 pp. N. Lloyd, London. - 1960. The phylogeny of Coleoptera. An1uta1 Rev. Ent., 5: 111-134. DALLwITz, M., T. A. PAINE& E. J. ZURcHER,2000a. 'Principlesof interactive keys'. http://biodiversity. uno.edu/delta/. & - 2000 b. User's Guide to the Delta Edi tor. Edi tion 1.03. CSIRO Di vision of Entomology, Canberra, 34 pp・ NewScirtidGenusNippot1oyphon fromJapan 527

FRIEDRICH, F., & R. G. BEUTEL,2006. ThepterothoracicskeletomuscularsystemofScirtoidea(Coleoptera: Polyphaga) and its implications for thehigh-level phylogeny of . J. Zoot. Syst. E、iel. Res., 44: 290- 315. HANNAPPEL, U., & H. F. PAULUS, 1987. Arbeiten zu einem phy1ogenetischen System de r H e1o didae (Coleoptera) - Feinstrukturuntersuchungen aneuropaischen Larven. Zoo1. Beitr., (N. F ), 31: 77-150. - & - 1991. Some undetermined He1odidae larvae from A ustral ia and New Zealand: fine structureofmouthpartsandphylogenetic position, pp 89-128. In: ZUNINo, M., X. BELL t s & M. BLAS (eds),AdvancesinColeoptero1ogy, A.E.C.,Barcelona. KAsAP, H., & R. A. CRowsoN, 1975. A comparativeanatomical study of Elateriformia and Dasc加oidea (Coleoptera). Trans r ent. Soc. Lolid., 126: 441-495. KUKALOvA-PEcK, J., & J. F. LAWRENCE,1993.Evolutionof thehindwinginColeoptera. Can. Ent., 125: 181-258. - & - 2004. Relationships among coleopteran su bo rde rs an d major neopteran lineages: Evidence from hind wing characters. Eu,. J. Ent., 101:95-144.. LAWRENCE, J. F., 1999. The Australian Ommatidae(Coleoptera: Archostemata): new species, larva and discussion of relationships. I,tvert. Taxi., 13:369-390. - 2001. A new genus of ValdivianScirtidae (Coleoptera) with commentsonScirtoidea and the beetle suborders. In MORIMOT0, K., K. MIZUN0, Y. HAYAsHI, T. Ito, N. ITO, K. AND0, M. TANIKAD0 &S. SHIYAKE,S. (eds),Sukunahikotla.Spec.Pub1.coleoptero1. Soc., Osaka, (1):351-361. -, N. B. NIKITSKY, & A. G. KIREJTsHUK, 1995. Phylogenetic position o「Decliniidae(Coleoptera: Scirtoidea) and comments on the classification of Elateriformia (senst1 late), It1 PAKALUK, J., & S. A. SLIPINSKI (eds),Biotogy, Phylogeny, andClassificationofColeoptera.PapersCelebrating the80t/1Birthday of Roy A. Crowson, 375-410. Muzeum i Instytut Zoo1ogii Polska Akademia Nauk, Warsaw. NIKITSKY, N. B., J. F. LAWRENCE, A. G. KIREJTSHUK & W. G. GRATsHEv, 1994. A new beetle family, Decl iniidae fam n., fromtheRussianFar East and itstaxonomic relationships(ColeopteraPolyphaga). Russ. Ent. J., 2: 3-10. NYHoLM, T.,1972. ZurMorphologicunci FunktiondosHelodiden-Aedoeagus(Col ).Ent.Scatld.,3:81-119. - 2000. Newspecies, taxonomic notes,andgenitaliaof NewZealandCypho1! (Coleoptera:Scirtidae). N. Z Ent., 22: 45-67. PIc,M.,1914.Dasci11idae, He1odidae,Eucinetidae.CoteopterorumCata1ogus,pars58.65pp.W.JUNK,Berlin. SAKAI, M., &M.SAT0,1996. Thecoleopteran family Decliniidae(Elateri「ormia,Scirtoidea) new toJapan, w i th description o f i ts second representative. Elytra, Tokyo, 24: 103-111. YosHIToM1, H.,2005.Systematic revisionof the familyScirtidaeofJapan, withphylogeny,morphology and bionomics(Insecta: Coleoptera,Scirtoidea).Jpn. J. Syst. Ent., Molt.Set・., (3),212pp.Matsuyama.