Phylogenetic studies of the subgenus Nialoe (s. lat.) (Coleoptera, Carabidae, genus Pterostichus), part 1: homology of the component parts of male genitalia and higher phylogeny

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Phylogenetic Studiesof the Subgenus Nialoe(s.Iat.()Coleoptera,

Carabidae ,Genus Pterostichus) ,Part 1: Homology of the
Component Parts Male Genitalia HigherPhylogeny
of and
K6jiSasakawa"

Graduate School ofAgricultural and Liie Sciences, The Universi otfy folqxo,Ytayo i1-1-1,
Bunkyo-ku, Tbkyo 1 13-865Z Japan
ABSTRACT-A phylogeneti chypothesis of the subgenus Niatoe (s .Iat .o)f genus Pterostichu sis proposed
based on a cladistic analysis of seventeen morphological characters. Through comparative studies of the
sclerites on the endophallus of ma[e genital iia tb,ecame apparent that the arrangement of these sc[erites
reflects seven differe onrtigins, The resuits show that Nialoe (s .Iat. )js monophyletic and composed of four
clades, while some tradition atlaxa are para- or polyphyleti ca,nd are defined only by symplesiomorphies.
Pterostichu s(Niato ten)osaicus sp. nov., which is indispensab lfeo rtracin gthe homology of sclerites in this
subgenus, is described.
Keywards:Niaioe (s ,[at, )s,clerite, homology, phylogeny n,ew species
INTRODUCTION Kitwsotnyp e(1sp9ec9i e6dse)scPr.itbeerd eKtoirsae,nadniBaelrole ofvroamn dSoPultuht eKnokroea(, 1w9it9h7)

The subgenus Nialoe Tanaka, 1958 is one of the more erected Natatianoe for P. rr]icmps of Wladiwstok, Russia,
diverse subgenera ot the genus Pterostichu sBonelli ,181O, which had formerly been referred to Paralianoe (Kryzha-
and is characterized by the sexual morphology of male ster- novskij et al. ,1995).
num 7 in most species. This subgenus is apterous, and its Thus, seven subgenera have been proposed to contain
distribut iios nprimaril ythe Sea of Japan region of the Far the 120 species fir sgtrouped int othe subgenus Nialoe.
East (Jedli6 1k9a6,2; Tanaka, 1985; Kasahara, 1988; Kryzh- However, their phylogenetic relationships have never been
anovskij etaL, 1995; Park and Paik, 2001; Bousquet, 2003). established on the basis of reliable evidence, and there is
The subgenus include sapproximately 120 species that considerable incongruenc eamong classifications by differ-
have either been united into one subgenus (Lianoe: ent authors, Jedlick a(1962 i)ncluded all Nialoe species in
Jedli6ka ,1962; Niaioe :Habu, 1981; Pterostichus :Tanaka, Lianoe .Habu (1981 c)riticized the superfluous split of sub-
1985) ,or separated int oseveral subgenera (Kasahara, genera, and treated them a[ 1as one subgenus, Niatoe.
1988; Park and Paik, 2001; Bousquet, 2003). Tanaka (1 985) ignored the subgenera erected for the Japa-
Niaioe was orjginally erected by Tanaka (1958 )for nese species and regarded them all as members of the sub-
Pterostichu sasymmetricus Bates, 1883 as a subgenus genus Pterostichu s(s ,lat. )L.ater ,Kasahara <1988 )dMded
related to Lianoe Gozis, 1882 of the Pyrenees. Tanaka 65 species into eight groups based on external structures
{1958 i)ncluded in the subgenus specjes that have an aede- {sexu aclharacteristics of male $ternum 7; number of setae
agus similar in shape to that of P. asymmetricus. Around the on antennal segment 2) and parts of the male genitali a(right
same time, lshida(1958)established Patelianoefor P. paramere;some of thescleriteson theendophal[us). How-

uenoi, and included it sallies in that subgenus. Later, ever, he did not discuss the phylogeneti crelationships
Nakane (1979 )proposed EPiniatoe for P. spiculiier. In the among the eight groups.
same paper ,Nakane and Straneo described a new subge- Taxonomic problem sarose in this group partl byecause
nus, Daisenialoe ,to accommodate P. ftij'imu rwahiic,h has no researchers had used male endophallic structures, which
some peculiariti eisn it sexternal structures, Kasahara and are known to be useful phylogeneti ccharacteristics in the
lto (1987 d)escribed Sphodroleronia, with it stype species P. Pterostichi n(iAlle 1n9,80; Koch, 1986; Nemoto, 1988, 1989a,
miyamai, emphasizing it sexternal peculiariti esP,ark and b; Brandmayr and Brandmayr ,1994; Bousquet ,1999;
" C o r re s p on d in ga ut ho r. PFhaox ne/: ++8811--33--55884411--75525257;; Sian ssoamkeaw Nai, a2l0o0e4 ,sp e2c0i0e5sa,, t hbe; pSreasseankcaew oaf asncdle rKiutebso otna , th20e 0s5u)r,-
E-mail/[email protected] face of the endophallus was known (Tanak a1,961; Nemoto,
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1988, 1989a ;Park and Kwon, 1996; Morita and Kanie, 1997; group ,rather than to adhere strictly to a forma ltaxonomy.
Morita ,1996a, 1998, 2001; Lafer et aL, 2001; Sugimura,
2o0n0l2y) ;i n hpoawretviecru, laa urtghrorosup su se(dT atnheamk aas1,9 d6i1a; gnKoa$staih cacrhaar,a ct1er9s88; SYSTEMATIC ACCOUNT
Park and Kwon, 1996; Morita ,1998), rather than examining Subgenus Nialoe Tanaka, 1958
their homologies among taxa. Nialoe Tanaka, 1958: 82. Type species: Pterostichu sasym-
The main purpose of the present study was to elucidate metricus Bates, 1883
the phylogeny of the subgenus Nialoe (s ,lat. ).First ,1 Nialoe as treated here includes East Asian species
describe a new species that has proven to be useful in trac- which Jedli6ka (1962 )treated as Lianoe; Habu (1981 )as
ing the homology of endophallic sclerites. Second, 1 review Nialoe ;and Kasahara (198 8as) Pterostichu s(s I.at, l)n, the
the homology of the component parts of the male genitalia, presen tstudy, the presenc eof setae on the ventral side of
primaril ythe sclerites on the surtace of the endophallus. the Ias ttarsa lsegment provide ssupport for monophyly of
Third ,1 propose a phylogeneti chypothesis of Nialoe (s .Iat.) this subgenus (se ephylogenetic section), The species
using cladistic methods and based on seventeen morpho- which 1 examined share the followin gcharacter states.
logica lcharacters (10 male genitali a,1 female genitali a6, Description [,Exter nstarluctures]/ Body medium to iarge
external structures}. (minimu m9,.5mm: P. (N. )ohkurai Morita ,[Mori t1a9,96b];
maximum, 24mm: P. (N. )ishikawai Nemoto [Nemoto,
1988]).Dorsalsurface of body blackand opaque toshiny

MATERIALS AND METHODS
without metallic ]uster .Legs reddish-brown to black ,Head
Most ot the specimens used in the present study were from the normai to large ;antennal segment 3 without pubescence
author's collection T.he rest of the material was loaned from the fol- except for P. ftij'irnurei iPa. nsdhotaroi, Pronotum cordate to
lowing publi cor privat ecollectiens /Laboratory of Forest Zoology, almost square, with single laterobasa limpression on each
DLNeiaptta eirSotcmnieaenSltncci eoest n ,TcFhoereMe usUstne Siucvmie,ernsTcie etko f,yyG oTr,oaJkdayupoaa,t nTe S{ockNhySooMo, TlJ )oaf;p NAaagntr i{icoFunZlatUlluTnrs)a ltT;anihdteute sniudme, 7E wliythtr saelxuoanlg- ocvhaarla c;tienritsetircval sinno tmo sitnt seprecriuesp.te d [M.aMlael esgteenr--

for Agro-Environment aSlciences ,Tsukuba, Japan (NIAES a)nd the italia ]A:edeagus robust; right paramere short to elongated;
collection of Mr. K. Akita (Mie J,apan) ,Mr. H. Karube (Kanagawa lef tone square, Endophallus large ,with vent toward lef tlat-
Prefectural Museum ef Natural History ,Kanagawa, Japan), and Mr. eroventral to ventral directio ne,xcept tor P. crpacipennis and
Kgd.ie rnMeeacrttsal u yma.Troaetxo oni onm(cilKcuadeg dia nwifnoa rJ,tmahapetasineo) n . coMonlo lsettcwt oioo fni 'nts,dh ie saptneydnp se awsbeplreeec ie sspeexc aimmoiefnn essd,ub- isct[s earloltiiezs ewdi thwi tshh gorotn, osptorariag hlpt ieencdoe p.[haFlleums;al gegeonniotpaolri e aMwee]adk:liyan
male P. tereti s(ty psepecies of Koreanialoe; femaie is unknown) ovjduct connected to semina- canal jus tanterior to the
and temale P. microps (ty psepecies of Natalianoe )wa,s obtained vagina; apophyses of basa lpar tof median oviduct and sem-
from the peer-reviewe dliterat u(rJeedii 619k6a2;, Park and Kwon, ina lcana[ weakly to distinct lsyclerotized; receptaculum
1996; Berlov and Plutenko, 1997>, Morphological characters not almost straighttostrongly bent,
referred to in the literatur weere coded as missing. Four spec[es, P.
(RP t(ehtryoossto i$d. c u$shtscrh)u.o) nschreims'tatus F{aDlduefromau nPrn. ),(a,Lndi aPn. o due(l)Mou. r}il a(cDuenojseusan), Pterostichus (IVia lmooseai)cus sp. nov. {Fig 1.)
roubaSianus (Lutshn iwker)e, used foroutgroup comparison, Dissec- This species is unique in having a mosaic combination
tjon technique$ and the terminology for female genitali afollew of external structures of some Korean and Japanese spe-
Bousquet (1999 )b,ut new terminolegy is proposed here in ter most cies. The sexual condition of male sternum 7 is consldered
othercharacters.
Seventeen characters (10 male genitalia ,t female genitalia as an apomorphic character state also possessed by P.
and 6 external structures )were observed from adults of 30 species, touzatin iand it sallies of the Korea Peninsula and the adja-
and all characters were treated as unordered (Append i1x). Phylo- cent islands .On the other hand, the opaque female elytra
geneti canalyses were performe dby PAUP' ver. 4.0 (Swofford, are considered as an apomorphic state shown in common
2cba0ot0te2sh) eoqwfui tahrla n daao nmdhe u tsruricesceetssis, c iswveieatrhch , TcBhR aTr ahbcetre arannac lywhs eessiwgahptpii nngbgeg a{ npFe arfrfroorrm im 1es19d0,6 9ur)se.pi lnigI-n w(itKha ssoameha r s1pae9,c8i8e)s. Hoofw etvhee rP,. tshpeh oenddropohfalolrusmis g orfo uthpi so fs pJecaiepsan
successive weighting, each character was weighted by the rescaled is a combination of presumed plesiomorphic character
consistency index (RC) ,but tree lengths, the consistency index (Ci}, states, whereby i tis possibi teo trace homologies between
awecn[vadtathleus a.tethqe ude aClrhle yatuer snwiatenicigogtnh et a eri dcnbhadoneo gxcte h$sa(trraRca lt pew )freposr,err to hbSear tebrcaeiostnlusili ttsrctautanycltitc ecu sdltuaapritpeeo derstu s wi fenrfrgeoo rm nAcoC1adC0lecT0suR lAa rNtweeapdsli- sKo am seaDh iaKarogarn eoaansndi s a .MnTadht isJsua mpsopatenocei,se es19 s9ip0s e cseiixemtsieir an(ra Fltloiy ,gP b.s2 u-,(t4 Ne)a.. s)igloyt odeinsstiisn-
optimization. guishe dfrom the latte rby the shape of the aedeagus and
My series ot taxenomic studies of this group treats all species the prominence on male sternum 7.
under the subgenus Niatoe (s .Iat, ,i,e ,,the subgenus Nialoe sensu Description. [Exter nstarluctures]/ Length: 8" ,18.8-18.9
Hitty apibes u n,so tp1e c9ai8els1w);a ;Nys em thpeeotrsoes,fi ob1rl9ee 8 ,t8toh, ee1x 9apm8hi9nyeal}o ,fg oaelr nlt ehstpe ei fcrcioeellsalt ioodwnisih rnirepgesca$tQ ln(psyr.ionp oFcsielrdustd,ingamnmd s(hnin=y,2 )e\x,,c e1pt9. 4f omrm fe m(an[=e 1e l)Dy.torras awhl iscuhr taacree oopfa qbuoed.y bLleagcsk
may change subtly in futur eanaiyses, Second, i tis importan tto reddish-brown. Head normal, Pronotum cordate; surface
enhance our understanding of the phylogenetic relationships of this smooth except for laterobasa limpressions which are slightly
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gp
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Fig. 1. Pterostichus <Aliat moosea)icus sp. nov. (A )Dorsal view. {B )Dorsa lview of endophallu$. (C )Ventral view of endophallus. (D )Right lat-
eral view of rjght paramere .gp, gonopora [piece ;rpb, rlght pigmented band; lpb ,lef ptigmented band; bs, basal sclerite; lpl ,[ef tpreapica llobe,
Scale lines :5 mm for (A) 1; mm for (B-D) T.he basal part of the aedeagus is omitted.
punctate. Elytr along-oval ;three dorsal pores on interva l3. tion,
Ventral side almost smooth; submentum with a pai rof setae. ParaCyoes. 1 841 9,same data as holotyp ei,n NSMT
Male sternum 7 with apical prominence ;prominenc ebroad collection.
and trapezoidal ;prominence narrower posteriori tyhan ante- Other specimens examined. Pterostichu sgotoensis
riorly; surface of prominenc eshallowly cup-shaped from the Kasahara and Matsumoto: 1 8, Wakamatsu-goe, Wakamat-
ventral view. [Male genitali aA]ed:eagus without a distinct sujima ls. ,Got6 lsls .N,agasaki, Japan, 1. v. 1988, (T ,Mat-
tubercle ,and shallowly dished on the lef tventral side near sumoto), hoiotype ,(NSMT> (endophal lnouts everted); 19,
ostium. Endophallus large ,moderately bent in lef tventral ditto ,18, viii, 1992, (K .Matsumoto), (NSMT),
directio n;pigmented bands present on both latera lsides Notes. Shape of spermatheca could not be observed
near ostium; sclerotization of the bands weak; basal sclerite because of damage during dissection.
distinc tw,ith swefiljng more on the dorsal side; right surface
of endophal]us swollen; sma]1 but distinc tswe[ling present
on dorsai surface near gonopore; weakly hooked lobe HOMOLOGY OF THE COMPONENT PARTS
present on right ventra] side near ostium; distinc tlobe OF THE MALE GENITALIA
present on lef tside near ostium; apical lobe weakly but dis- ln the present study, seven sclerites of differen torigins
tinct lsyclerotized; gonopore weakly sclerotized, with gonop- are recognized on the endophallus, the homology of which
oral piece, Right paramere short and rounded apically. Left are considered as follows.
paramere square, IFemal egenitalia ]V:agina with slightly
sclerotized part around median oviduct; surface without con- Right pigmented band (rp bin Figs ,1, 2; character 2 in
spicuous pigmentation ;apophyses of median oviduct and Appendix 2)
seminal canal weakly sclerotized over their entirety. ln some Nialoe species (e,g .P,. (N. )mosaicus), there
Locality .Fukuejima [s. ,Got6 lsls .N,agasaki, Japan. is a pigmented band on the right laterodor ssaurlface of the
Holotype. 8" ,Mt. Sasadake, Fukuejima ls, ,Got6 lsls., membranous part near the ostium. The pigmentation varjes
Nagasaki, Japan; 1. x. 1996, (H ,Karube), in NSMT collec- from rudimentary to distinc att the subspecific Ievel.
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Fig. 2. Homology of scierites on the endophallus. {A }Pterosnchus (tVia ttsohiezu}chiensis <B )P. <N. s)ymmetncus (C )P. (N) mosatcus (D)
P. (N, m)ucronatus, (E >P. (N. a)bacilorrnis (F )P. (N. c)tandonis (G )P. (N) katashinens inasganoensts, (H )P, (Liano edu)lou" .rpb, right pig-
mented band; lpb ,lef tpigmented band, bs, basal sciente; mpl, median pigmented lobe; lpl ,Pet tpreapicaHobe, as, apical sc]ente Soli dlines
referto homology of the basal sc[erotizatton (bs )a,nd dotted ones to that of the lef tpreapical lobe 1ndicatio onf$ the gonoporal piece are omit-
ted because ot their obvEous homo]ogy. Scale 1ines ,1 mm
Left pigmented band (lp bin Figs .t-4; character 3 in Tanaka, 1961, "median macula" sensu Nemoto, 1989a, part
Appendix 2) of the "pigmented pad" sensu Nemoto, 1989a, and "apical
There is a pigmented band on the lef tlatero -to latero- copulatory piece" sensu Monta, 1998, ln Tanaka's (1961)
dorsa lsurface of the membranous part near the ostium. The study, the endophallus was not everted, so references to
pigmentation varies from rudimentary to distinc att the sub- "apicai" and "basal'' in the positio nof these sclerites are
specific level .In R (N. )spicubterand it sallies (= EPiniaioe reversed Nemoto (1989a n)amed th[s character "median
Nakane, 1979 sensu Kasahara, 1988), the posterio rend of macula" However, most species have the sclerite at the
the pLgmented band is connected to the anterior end of the basal par tof the endophallus, so that [`median" is unique in
basal sclerite. This character is synonymous with "poorly P. (N. )piesiomotphu swi,th which Nemoto <1989 ade)fined
sclerotized plate "sensu Morita and Kanie, 1997 However, this structure's terminology (Nemot o1,989a: 3, Fi"g m,e3d). jaInn
their observation was based on specimens where the the same paper ,Nemoto (1989a c)onsidered the
endophallus was not everted (Mori tanad Kanie, 1997: 166, macula'' (bas ascllerite in the presen tstudy) of P. (N. m)ucr-
Fig .3), so that only the basal part of this band was onatus and P. (N) shiibanus to be a ''pigmented pad' '(left
observed. Therefore, "plate" is an inappropnate term tor th-s preapic allobe in the presen tstudy; Nemoto 1989a: 4, Figs.
character 9-12) These two species do have one sclerite on the left
latera lside on the endophallus; however, the position sof
Basa] sclerite (bs in Figs. 1-4; character 4 in Appendix 2) the sclerites are not correctly shown in Nemoto's (1989a)
There is a sclente on the let tlaterodor stao llatera lsur- illustratLons because the endophallus was not ful[ yeverted.
iace of the membranous part near the ostium. It sshape and 1 examined the completely everted endophallus of P. (N.)
degree of sclerotization vary at the subspecific level ,Th:s mucronatus and P. (N. i)shizuchiensis (probab al lyoca lrace
character is synonymous with "apical sclerite" sensu of R {N. )shiibanus), and confirmed that these species have
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Fig. 3. Endophall oif NiaJoe (s ,lat .(>rig hlatter avliew, let ltater avliew), (A )Pterostich u(sNialo feL)ijimu r<aBi. )P. (N. )min'ficus, (C )P. (N,)
yakushimanus. (D }P. (N) piesiornorphus. (E )P. {N. )miyamai. (F )P. (N. )rnacrogenys. (G )R rnicrops. (H )P. (rw y.o)shiclai .(1 R) {N. b)eOator.
(J )P, (Al .op)acipennis, rpb, right pigmented band; lpb ,lef tpigmented band; bs, basal sclerit e;mpl, median pigmented lobe; lpl ,lef tpreapical
bbe. Indicatio onfs the gonoperai piece are omitted because of thei robvieus homology, Scale "nes :t mm,
the sclerite that Nemoto (1989 at)ermed the "pigmented Median pigmented tobe (mpl in Figs. 2, 4; character 5 in
pad" (le pfrteapic allobe in the presen tstudy), but at a more Appendix 2)
basal positio non the endophallus than in his illustration, There is a weakly to strongly pigmented lobe on the
Thus, because Nemoto <1"m9e8d9i aamni)sidentified the homology middle dorsa lsurface of the endophallus in some Nialoe
among the sclerites, the macula"' sensu Nemoto, species (e.g P.. ,(N. s)ymmetricus) and outgroup species (P.
1989a is not always "macula". Morit a(199 8i)llustra ttheids (Liano ed)ufour aind P. (Pterosti cs.h sutsr. )cristatus) .This
character of" aPp. i(catlV a.ba)ciicrmis and it sallied species and character is a synonym of "median lobe'' sensu Nemoto,
named i t copulatory piece ".However, as with 1989a. The detiniti oonf this character is exactly the same
Tanaka' s(1961 s}tudy, "apical" i san inappropri atteerm for as that of "medjan lobe "in Nemoto (19B9 ai;n P. (N. )mucr-
this character. onatus; however, i twas overlooked, presumably because of
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Fig. 4. Endophalli ot Nialoe (s .Iat.) ,Ptetostk:hus (s s.tr.} and Axlyosodus (rig hlattera lview, lef tlatera lview). (A )Pterostichus (Nialo mea)cro-
cephatus. (B }P. (N. s)phodrolorrnis. (C )P. (At m.as)hidai. (D )P. (rw s.in)cerus. (E }P. <tV p.r)aedo. (F )P. (N. u)enoi. (G )P. (N. o)rnogoensis. (H)
P. (N. l)atisty i(i1 sP.). (Pterost is.c shtru.) scristatus. (J )P. (dyosod ulsac)unosus roubalianus. rpb, right pigmented band; lpb ,lef tpigmented
band; bs, basal sclerite; mpl, media pigmented lobe; ipl l,ef tpreapical lebe ;as, apical sclerite, lndicatio onfs the gonopora lpiece are omitted
because of it sobviou$ homology. Scale lines /1 mm.
insufficie nevtersion of the endophallus [p4 ,Fig, 11]) .In the i tis djstinct lscylerotized and concave (e.g P.., (N. >symme-
present study, 1 propose the term median pigmented lobe for tricus) ,By tracing homologies among species, i tis reveaied
this character, because the term "median lobe" means that this character is a specialized form of the lef tpreapical
aedeagus or penis in the general terminology of the lobe <Fi g2,). This character is synonymous with "lobe a"
Coleoptera, sensu Nemoto, 1988, part of the "pigmented pad" sensu
Nemoto, 1989a, and "apical copulatory piece' 'sensu Morita,
Leftpreapicallobe{lpilnFigs.1-4;characters 6and 7in 2001.

Appendix 2)

l nmost Nialo especie$, the dorsal surface of the left Gonoporal piece (gp in Fig. 1; character 8 in Appendix 2}
preapical lobe is more-or-less sclerotized, ln some species, AII species of Niaioe have a weakly sclerotized gonop-
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ore, of which the dorsal aspect is strongly sclerotized. Many was not recovered in the consensus tree (Fi g5)..
authors have deseribed the dorsal scletotized portio ans a From the successive-weighting analysis, fiv etrees were
"copulatory piece" or used an analogous term ("api cscalelr- obtained, Their tree lengths, Cl, and Rl were equal to those
ite "[Tanak a1,961] ,"copu[atory piece "[Mori t2a0,03] "prox- of the equal-weighting analysis. Tree resolution was better,
imal copulatory piece "[Mori 2t0a01,]). Only Nemoto (1988, and four lineages were emerged (Fi g5,, 6) .The monophyly
1989a) distinguishe di tfrom other sclerites on the endophal- of the Pterostichus (s .str.> + Lianoe + Nialoe (s ,lat. l)ineage
lus ,and named i tthe gonopora lpiece .]n the present study, was supported by a moderate bootstrap value (78%).
1 adopt Nemoto:s (198 819,89a) term for this character. Although the bootstr aspupport is re]atively low (57%} t,he
monophyly of tViaioe (s .Iat .w)as supported by the absence
Apical sclerite {as in Figs .2, 4; character 9 in Appendix 2) of setae on the ventral side of the las ttarsa lsegment {char-
ln some Nialoe species (e. gP.. ,(N. )katashinensis acter 14), This character state is a synapomorphy of IVialoe.
naganoensis>, the ventral surface of the endophallus near Lineage A comprises the species that Kasahara < 1988)
the gonopore i srudimentarily pigmented .Some outgroup referred to as the tnirificus ,ftijimu raasyimm,etricus, spiculi-
species (P .(Liano deu)touriand P. (Pterosti cs.h sutrs,) cris- iec and uenoi groups .The shape of the endophallus (char-
tatus )have a large sclerotized projecti oinn the correspond- acter 1O) and the presenc eof setae on antennal segment 2
ing spot, so 1 propos ethe term apical sclerite to represent <charac t1e2r) support the monophyly of this lineage ,Lin-
this character. No other authors have described thi scharac- eage B includes P. (N .ma)crogenys, P. (N. b>allato rP,. (N.)
ter, opacipennis, and P. (N. )tereti sa,nd it smonophyly is sup-
ported by pigmentation of the innermost part of the ventral
PHYLOGENY AND CHARACTER EVOLUTION s(idNe. )moifcr otphse, va gPi.n a(N (. pc)rhaeadro aan cd1t1 P)e., rL(iNn.e a)signec eCru sc.om pTrhisees mono- P.
From the equal-weighting analysis, 20 trees of 70 steps phyly of this lineag ei ssupported by the opaque dorsal sur-
were obtained, with Cl=O.586 and Rl=O.729, The strict con- face of the pronotum and elytra <chara c1t3)e. rThe remain-
sensus tree was we]1 resolved at the species level ,except ing species are included in Lineage D, monophyly of which
fo rLineage A (defin beedlow), where monophyly and some is supported by the presence of the basal sclerite (character
interna lnodes were not supported. The monophyly ot Niatoe 4). In all lineages (A-D )mo,nophyly and most of the species
Kasahara(IY88)
'l']iesl'minetric"sgroup(sllbgenusnotrnentioned)
60.VJ,MMets'ir''us
totEalini'k),afoushinianus'
Thcvakuslti}nanusgroup(s"bgenusmotmentloned)
"''"'"1/....The(veaL'ipennisspccicsgroup
mac'ro['ephtTiw,y
piesiomony.ihu,y- r･ltThecx,5ac'ifi)Fmissp:ciesgraup

50inucT'oanhalctM'f/or.mi/s'
S2･
nio,saiL'lts#v'oshiditi
････-･･･････

;is'ii]7･･l:t tZThse2 sh1iiba:nrfs species group

LineageD il'i"'i lTh'c's'f)lh･o'tt'i''ioft)i'misspeciesgroup
sphoth't?foJ'mi･s 'i'henn/vamaispccirsgroup
:2lh I.t..Ii.I.r:. .dI./ :a.-l :.C;t.su:./b.2c.]Ii.Tu.sii.Y-?hodrofb (is' o.stnr.i)a}
Sin(/e/'1.iSSpJ'c)edoStnier()s.'/,ste/etiseopacipennis'
LineageC ''I'he
t)/t'[cJ'ogen ".'/ specics gruupThesphodrtijbrmisgroupCsubgenusnotmentielLed)
M'ttioe(s.Iat.) LineHgeB hetiatoJ' macirose)1),S
･･-･･･-･･･
5.7
.tiftisltzil-cri 1Theflijimttr"igro"pCisubge]vsDaisenittloe(s.stn))
kEtslsinensi.vnaga"oen,vis
al,ha'Jt'Mimasti''iAc',lilsisdandonis,spicuTTihhieeflecaj"st',)'J,ii,TmsaieCttrysic'uusieinsssipseb'cpeicieessgrogu.proup ] he "wmmetricus group (= s.ubgeT]us vtaiee {s stn})
7S 1Thespicuiijergrotip{=subgenusopintatoe(s.sdn))
L uenoi Thetfftnoispecicsgroup 1'i
On70gOeSISisnTir(lfic'ifsc'rwfbziTi'hieCtS'simS'tCxIftgUStslcv'haoinnheasnvH'isspeciesgroup hc ltenoi group (= subgenus paraii"noe cs str,))
The mir{'fieus group (subg¢nus not mentioneti)
9S
outgl'oup
[

i'oifbaiitznus
Fig .5.suSstric ctonsensus tree of the 5 trees obtained by the succesive-weighting method. Closed circles indicate nedes shared by the consen-
trees o f t h e 2 0 t re e s re su l ti ng f r o m th e e qu a l- we i gh ti n g an al y si s The taxonemy from Kasahara ( 1988) is represented fer comparison, Spe-
cies with an asterisk are not include din Kasahara' s(1988 s)tudy.
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12t2 K. Sasakawa
, 1
O i-2
zts
tensis

1
Fig. 6. Phylogeny and possibl echaracter evolution of Nialo e(s.lat. )based on one of most parsimonious trees obtained by the $uccesive-
weighting method. Numbers in the squares indicat echaracters in Appendix 2, and numbers above the squares indicate changes in character
states. Character-sta ctheanges were reconstructed with ACCTRAN optimization,
relationships werenot supported by high bootstrap values. eny of Nialoe was full yresolved, except for re[ationships
among the four larges tclades (Linea gAe-Ds). The trees
DISCUSSION obtained here suggest that some tradition atlaxa are not
monophyletic.
This study provides for the firs ttjme a phylogenetic In Lineage A, monophyly of each of the groups pro-
hypothesis of Nialoe, based on the cladistic analysis of 17 posed by Kasahara (198 8i)s supported, except for the
morphological characters. I t became apparent that asymmetricus group. The asymmetricus group is paraphyl-
endophallic structures are wejl developed and diverged etic i nall trees obtained, because the tLtiimuraigrou pcon-
within Niatoe, and that they can contribute significantly to the sistently occupies the most derived branch of the asymtne-
reconstruction of the phylogen yof thi sgroup, The phylog- tricus group + ftijimur agiroup iineage .The characters that
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Phylogeneti Sctudie soiNialoe Part1 1213
Kasahara (198 8r)egarded as diagnosti cof the asymmetri- 34/1-29
cus group, (sexu ach]aracteristics of male sternum 7, more Berlov O, Plutenko (1997 T)wo new subgenera of the genus Pteros-
trhiagnht vtehnrterea ls estuarefa coen oafn ttehnen aaled esaeggumesn,t a n2d, tau bsheorrctl e too ne ltonh-e VAtekisactdhneu ims{ kiCli ori5kl: ue4t7os-pk5t o1Gey orCsaaur,daabrisdtave e)ninroo ym Stehle' sFkaor kEhaoszty aoty sRtuesvsejnan.oy
gate right paramere )are a[so present in the ftijimuraigroup. Bousquet Y (199 9S)upraspecif icclassification ot the Nearctic
In other words, these are symplesiomorphies of the asym- Pterostich i(niColeopt eCarraa:bidae ).Association des Ento-
metricus group + ftijimuraigrou plineage. mologists Amateurs du Quebec, Quebec
th e sIpnh oLdrionioeramigses B- Dg,r omounpop hywlays n ootf ssoumpeport esdpecie s(F gir go5.u)p s. Aosf B o u sqoafgua e-tAPda eYplh aa(gea2a"r0c0t3 iT) cCrEiodbl eeb oyP pt1et rLeoOrs batVli oc,Alh uiSmn meiBleotn aeinAlraic, ,h1Ao8ps1otOje,[m oal tBna o-"oCMaktaylsoxg,ueo pSht-en-
stated by Kasahara <i988 )th,e sphodroicrmis group is less strup, pp 469-520
specia]ized externally, except for the two macrocephalic Bandmayr P, Brandmayr TZ (1994 >The evolu"tiCoanarryab ihdistory ot the
species groups (th eopacipennis and macrogenys species genus Abax (Coleopte Craar,abidae) I.n beetles: ecol-
gthreo u"simpilasrity) "t;here foof rte h,te heexit errtnaalxon cohmaryac therass. been based only on Fa r roMgi LyJs S L aun<fdt1 , 9Je6v-oP9l uAM)t aisoeuncl"cfe saEsii dvtS e,pbry iKanp pgDreoe xrspi,mepa nt1ido9en-sr2 ,4M aDpupfroraecnhe, Mto Lchoarreaac-u,
The two macrocephalic species groups are both split ter weighting. Syst Zool 18/ 374385
within Lineages B-D {opacipen nspiecsies group: P. (N.) Habu A ( 1981) Female genital iofa Pterostich ispneicies mainly from
opacipennis and R (N. )betlatorare in Lineage B, P. (N.) Japan (1 1()Coleopte Craar,abidae) E.nt Rev Jpn 36/ 33-53
m(acNro, cme)apchraolguesnys i n iLs iinn eLaignee aDg; em aBcr,o gRen ys(N. )mi scrpoepcsies i ng rLoiunpe:a gPe. l s h iPEdt naeHt r Roes(tv1i 9cJhp5un8 s )S9B:o o5mn-ee8l lni ew(C eorl leiotp tlkt eneCoarwranab ,jsdpaecei)es fr oomf tJhaep agne n(us1),
Cb,e rasn dof P .e a(chIV y.gor)sohuipd aariei n Lsiimnielaagr et oD )e, alc htis tortuhee rt haitn thhae vmiemn- ga J e d lsiie6nk aA( (P1t9e6r 2oMo}sntog irTcarphhiiigeonn iod,teso m TirM,iybauds iP) te(rCosotlicehion piau ts-e OCrsaatraa--
large-size dhead, small eyes, and long mandibles, but the bidae), Abh Ber Staat Mus Tierk Dresden 26/ 177-346
pnorte smeonnot phrye]seutlitc stro ngalnyd stuhggaets tsthei t rhmaactr oecaecphhal iscpecies f ogrrmosu pa ries K a s atshipeacchriiaa nleS c ar(reaf1beir9den8c 8e b)De ietsto tltreh iesiibn r uJotra iipgpoiaannnt t. aen1drn n " Thasenpdec i bastepieeotcnil'a'ets i oEonfd J baoypf a Mnp, tS eawirttoOhs,-
convergent. T6kai Unive rsity Press ,Tokyo, pp52-65 [i nJapanese]
The evolutionary polariti esof the characters inferred Kasahara S, lt oY (1987 A) new Pterostich u(sColeopte Craar,a-
from the present analysis indicat ethat the diagnosti cchar- bidae )from the upper hypogean zone of central Shikoku,
aucatle rsdi moof rtphhei sspmho,d roonleorm isseta osnpec iaenstenn aglrou ps e(gmrenetdu c2t, iofaon dsnex -a K e c hs" noigur(ittdh'1w9es8t 6 <)M CoJorapip h.aCon,lro, agKbio.cn) atl,a-y cp0 oh5my5psl: eix1ol3o9 g-oif1c 4as il5bsltiundgi esspe coines P ltne ℃toasrtabiicdhus
short right paramere of which the apex is rounded) are sym- beetles t,heir adaptations and dynamics' E'd by PJ den Boer,
plesiomorphies of Nialoe. The polyphyly of the sphodroicr- ML Luff ,D Mossakowski ,F Weber, Gustav Fischer N,ew York,
mis species group resu[ts from it sdefiniti obny only synple- pp 269-279
KryzhanovskiOjL,BelousovIA,KabakH,KataevBM, MakarovKV,
siomorphies,
Shilenko vVG { 1995) A Checklis ott the Ground-Beetle osf Rus-
sia and Adjacent Lands (]nsec tCaoP,eoptera, Carabidae )P.en-
ACKNOWLEDGMENTS seft Publishers, Sotia
Lafer GS, Park JK, Paik JC (2001 A) new pterostichine beetle of the
1 thank Dr. KOhei Kubota, Assoc. Professor of The University subgenus NiaJoe Tanaka from South Korea (Coleopte Craar,a-
ot Tokyo, Tokyo, Japan, for his guidanc eand encouragement in the bidae). Insecta Koreana 18: 243-248
course oi thi sstudy, 1 also thank the tollewi npgeople for offering Morita S (1992 }A new subgenus and species of Pterostichus
specimens or cooperaiion in various ways/ Mr. Katsumi Akita (Mie}, (Coleopte Craara,bidae) from Aomori Preiecture, north Japan.
Mtro.r yH ,aKraunkaig aKwaar)u,be D r(, KRay6nsaugkeaw aIPsrheitkeacwtau r(a TlMoukseyuom )Mr, , ofT aNdaatsuhria l MaHei-s- Mo r itEal ySt r(a 2109/ 9156 -aA19 )litt lkenewn pterostichi cnaerabid, Pterostichus
hara {The University ef Tekyo), Mr. Shunsaku Mano (Tokushima), hozumii lshida (Coleopte Craar,abidae), Japanese Journal of
DMrr., MKuenieitcoh slMhait sMuamroutyoam a{K a(gTahwe Naa )tD,iro, nKael iSscuikeen cNeem oMtuoseu {m,Th eTUonkiyvoe)r,- Mo r itSay sSt e(ma1ti9c9 E6n btPot)meorloosgtyic h2u/s 1o3h3ku-r1a3i6 (Coleopt Cearraab,idae) ,a
s[ty ot Tokyo), Dr. ShOhei Nomura (The National Science Museum, new relative of Pterostichus latisly flriosm the subalpine zone of
Tokyo), Mr. Jun Ogawa (The University oi Tokyo), Mr, SOsuke the Japanese Alps, EIytra 24: 197-202
Shiokawa (Toky oD)r,. Teij iSota (Kyo tUoniversit y)M,r. Hiroshi Morita S (1998 A) new Pterostichus (Coleopte Craar,abidae )from
Sugaya (Hokka iUdnoiversit yD)r,. Yasuoki Takami (Kyot Uoniver- the southern Japanese Alps ,Elytr a26/ 297-302
sity), Mr, Mitsuyukl Wakabayash[ (Chiba) ,Mr. Osamu Yamaji Morit aS (2001 )A new Pterostich u(sColeopte rfaro)m western
(Okayama >D,r. KOj iYasuda (Natio nlanlstitu tfeor Agro-Environ- Kyushu, west Japan, Japanese Jeurna lof Systemati cEntomol-
mentaL Sciences ,Tsukuba) and Dr. Shin-ic hYioshimatsu (National ogy 7/ 35-39
]nstitu ftoer Agre-Environment aSlciences ,Tsukuba), Morit aS (2002 P)terostichj cnaerabid beetle sot the subgenus Cr)to-
bius (Coleopt Cearraab,idae )from north Japan, Elytr a30: 73-
89Morita
REFERENCES S (2003 )Two new Pterosik;h u(sColeopte rCaar,abidae)
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TThheZeo olZoogiocalloSogciiectyal Society ooff JJaapapnan
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167Nakane Sasakawa K {2005b P)terostich umsacrogen)/s Bates, 1883 (Cole-
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Bulleti not the Biogeographical Society of Japan 43: 39-45 Sugimura A (2002 A) new subspecies of Pterostichus tocia iMerita
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Bulleti nof the Biogeographical Seciet yof Japan 44/ 123-125 K (1958 )A new subgenus and two new specie$ of the
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(Coleopte rHaar:palidae). Korean Journal oi Entomelogy. 26/ tyO 261 78-83
93-103 Tanaka K (1961 )Studie son the genus Pterostichus from Japan
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ef Korea 12, lnsec tKoreana Suppl. 19, National lnstitut eof 49Tanaka
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Pterostichus Bonelli from Korea (Coleepte rHaar:palidae) 1. [Plat 2e0s-25] .Hoikusha ,Osaka [i Jnapanese]
MicroniaJoe subgen. nov, Korean J Entomology 26: 72-77 Tanaka K, lshid aH (1972 S}tudie son the genus Pterostichus from
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Bates, 1883 {Coleopte Craara,bidae). Biogeography 6: 69-73
Sasakawa K (2005 a)Taxonomic notes on Alyas Strum ,1826 (Receiv eMdarch 24, 2005 f Accepted September 13, 2005}
(Coleopte Craar,abidae, Pterostichini o)f Japan. Biogeography
7: 11-20
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