Arma custos ( Fabricius, 1794 )

Zhao, Qing, Wei, Jiufeng, Bu, Wenjun, Liu, Guoqing & Zhang, Hufang, 2018, Synonymize Arma chinensis as Arma custos based on morphological, molecular and geographical data, Zootaxa 4455 (1), pp. 161-176: 164-169

publication ID

publication LSID

persistent identifier

treatment provided by


scientific name

Arma custos ( Fabricius, 1794 )


Arma custos ( Fabricius, 1794) 

Cimex custos Fabricius, 1794: 94  . Syntype (s): [ Germany:] “ Kiliae ” [= Kiel]; ZMUCAbout ZMUC. 

Pentatoma custos: Latreille 1804: 188  .

Arma custos: Hahn 1832:95  .

Asopus custos: Burmeister 1835:379  .

Stiretrus custos: Blanchard 1840: 153  .

Arma custos var. fuscoantennata  [as fusco-antennata] Motschulsky, 1860: 501. Nomen nudum.

Auriga custos: Kirkaldy 1909a: 15  .

Arma custos  f. scutellaris Stichel 1961a: 656. Unavailable name (International Code of Zoological Nomenclature, 4th ed., 1999, Articles 10.2, 15.2: proposed as infrasubspecific). No types fixed.

Arma neocustos Ahmad & Önder, 1990: 8  . Holotype (♂): Turkey: Lüleburgaz; EUIT. Synonymized by Thomas 1994: 165.

Arma neoinsperata Ahmad & Önder, 1990: 9  . Holotype (♂): Turkey: Lüleburgaz; EUIT. Synonymized by Thomas 1994: 165.

Arma chinensis Fallou, 1881: 340  . Syntype (s): China: “ Pékin ” [= Beijing]; MNHNAbout MNHN. New synonym.

Arma discors Jakovlev, 1902: 64  . Lectotype ( Derzhansky et al. 2002: 367) (♀): Mongolia: “Khalkha-Gol” [= Khalkhyn gol River]; ZMAS. Synonymized by Kiritshenko 1922: 14.

Auriga discors: Kirkaldy 1909a: 15  .

Auriga chinensis: Kirkaldy 1909a: 15  .

Auriga peipingensis Yang, 1933: 21  , 44. Holotype (♀): China: “ Peiping ” [= Beijing]; lost. Synonymized by Zheng 1981: 29 (tentatively). 

Diagnosis. BOdy Oval, MEdIUM sIzEd, dUll brOwNIsh, rElaTIvEly flaTTENEd; hEad wIThOUT ElONgaTE pOsTOcUlar rEgION, ThErEfOrE EyE adjacENT TO aNTErIOr MargIN Of prONOTUM; MaNdIbUlar plaTEs NOTAbout NOT cONvErgENT apIcally; rOsTrUM rEachINg METacOXaE; aNTENNal sEgMENT II lONgEsT, bUT a lITTlE shOrTEr ThaN sEgMENT III aNd IV cOMbINEd. LaTEral MargIN Of prONOTUM sErraTE aNTErIOrly, aNd ThE OUTEr MargIN wIThOUT ObvIOUs black bElT; hUMEral aNglE frOM ObTUsE TO aNgUlaTE, dIsTINcTly sUrpassINg cOsTal MargIN Of fOrE wINg IN rEsT; ThE METaThOracIc scENT glaNd OsTIOlE wITh wElldEvElOpEd, ElONgaTE pErITrEME aNd a sMall black spOT sITUaTEd aT ThE MEdIal sIdE; prOfEMOra UNarMEd; prOTIbIaE NOTAbout NOT dIlaTEd. BasE Of abdOMEN wIThOUT TUbErclE. ( FIg. 2 View Figure )

Description. Body Of MEdIUM sIzE, brOwN aNd Oval, dOrsUM raThEr flaT, vENTEr MOdEraTEly cONvEX. Body surface and vestiture. BOdy vIrTUally glabrOUs, wITh shOrT, INcONspIcUOUs pIlOsITy ON aNTENNal sEgMENTs III–IV aNd ON lEgs; ThE whOlE dOrsUM wITh lOTs Of UNIfOrM dEEp pUNcTUrEs, aNd ThE vENTEr alsO wITh pUNcTUrEs, bUT ThE laTEral sIdE a lITTlE dENsE ThaN cENTEr

Head. FlaT, NEarly as brOad as Or slIghTly brOadEr ThaN ITs lENgTh alONg MIdlINE; laTEral MargIN wEakly cONcavE; aNTErIOr Of EyE, raThEr sharply EXplaNaTE, lEavINg aNTENNIfErOUs TUbErclE slIghTly EXpOsEd IN dOrsal vIEw; clypEUs slIghTly shOrTEr ThaN MaNdIbUlar plaTEs, apIcal pOrTION a lITTlE flaTTENEd; MaNdIbUlar plaTEs brOadly rOUNdEd aNTErIOrly, NOTAbout NOT cONvErgENT apIcally; bUccUlaE sIMplE, UNarMEd; EyE glObOsE; OccElI vEry sMall, sITUaTEd dIsTINcTly IN pOsTErIOr MargIN Of EyE, INTErOcEllar dIsTaNcE abOUT TwO TIMEs as lONg as dIsTaNcE Of ONE OcEllUs frOM EyE. ANTENNal sEgMENT I shOrT, apEX NOTAbout NOT rEachINg apEX Of hEad; sEgMENT II lONgEsT, abOUT 2.0 TIMEs as lONg as sEgMENT III. ROsTrUM rEachINg METacOXaE; sEgMENT I NEarly rEachINg aNTErIOr MargIN Of prOsTErNUM; sEgMENT II lONgEsT, shOrTEr ThaN sEgMENT III aNd IV cOMbINEd; sEgMENT III lONgEr ThaN IV.

Pronotum. SUbhEXagONal, abOUT TwO TIMEs as brOad as lONg; dIsk raThEr flaT; aNTErIOr MargIN dEEply cONcavE; aNTErOlaTEral aNglE TrUNcaTEd Or bEarINg a MINUTE, aNTErOlaTErally dIrEcTEd dENTIclE; laTEral MargIN a lITTlE cONcavE, fINEly sErraTE aNTErIOrly, ElEvaTEd pOsTErIOrly; hUMEral aNglE frOM ObTUsE TO aNgUlaTE, dIsTINcTly sUrpassINg cOsTal MargIN Of fOrE wINg IN rEsT ( FIg. 3 View Figure ); pOsTErOlaTEral aNglE acUTE. Scutellum. ElONgaTE, dIsTINcTly lONgEr ThaN ITs basal wIdTh; wEakly cONvEX IN laTEral vIEw; wITh 1+1 sMall IMprEssIONs aT ITs basal aNglEs; pOsTErIOr NarrOwEd pOrTION rElaTIvEly lONg aNd NarrOw, gradUally TapErINg TOwards apEX, rOUNdEd apIcally. Thoracic pleura and sterna. PrOEpIsTErNUM sIMplE; METaThOracIc scENT glaNd OsTIOlE assOcIaTEd wITh aN ElONgaTE, wEakly cUrvEd pErITrEME, a sMall black spOT sITUaTEd aT ThE MEdIal sIdE; EvapOraTOrIUM NarrOwly EXTENdINg TO pOsTErIOr pOrTION Of MEsIplEUrITE. MEsOsTErNUM dIsTINcTly carINaTE alONg MIdlINE, METasTErNUM slIghTly ElEvaTEd aNd flaTTENEd. Fore wing. COrIUM far sUrpassINg apEX Of scUTEllUM pOsTErIOrly IN rEsT, brOadly lEavINg cONNEXIvUM EXpOsEd; MEMbraNE dIsTINcTly sUrpassINg apEX Of abdOMEN. Legs wIThOUT pEcUlarITIEs; fEMOra UNarMEd; TIbIaE sIMplE, NOTAbout NOT dIlaTEd, wITh a brOad lONgITUdINal fUrrOw dOrsally.

Pregenital abdomen Oval; sTErNITE III wIThOUT a MEdIaN TUbErclE basally; pOsTErOlaTEral aNglEs Of cONNEXIval sEgMENTs wEakly prOdUcEd; sTErNal glaNdUlar paTchEs lackINg.

External male genitalia ( FIg. 4 View Figure ). GENITal capsUlE TraNsvErsEly Oval IN pOsTErIOr vIEw, pOsTErOlaTEral aNglEs (pla) rOUNd ( FIg. 4a View Figure ) rOUNd, sITUaTEd slIghTly hIghEr ThaN NOTch IN vENTrOpOsTErIOr rIM (vpr) ( FIg. 4b View Figure ); dOrsOpOsTErIOr rIM (dpr) cONcavE ( FIg. 4a View Figure ), wITh a paIr Of dOrsal sclErITEs (dsIM) ( FIg. 4a View Figure ). AEdEagUs sIMplE, wITh ONE paIr Of vENTrOlaTEral cONjUNcTIval lObEs (vlcl) ( FIg. 4c View Figure ) whIch arE OblONg, apIcEs arE sclErOTIzEd aNd NOTAbout NOT bIfUrcaTE; apIcal cONjUNcTIval lObE (acl) ( FIg. 4c View Figure ) sEMIcIrcUlar aNd NOTAbout NOT bIfUrcaTE. MEdIal pENIal plaTEs (Mpp) ( FIg. 4c View Figure ) sIMplE, ElONgaTE aNd sTrONgly sclErOTIzEd, laTErally flaTTENEd, pOsTErIOrly brOadly rOUNdEd. VEsIca ( FIg. 4d View Figure ) sTrETchEd OUTAbout OUT frOM ThE vENTEr Of MEdIal pENIal plaTEs, NOTAbout NOT rEachINg half Of MEdIaN pENIal plaTEs. ParaMErE shOrT aNd wIdE, wITh a shOrT fINgEr-lIkE INNEr braNch (fIb) ( FIg. 4E View Figure ), prOTrUdINg TO ONE sIdE.

External female genitalia. LaTErOTErgITEs VIII ( FIg. 5 View Figure : lT8) brOadly rOUNdEd, spIraclE VIII EXpOsEd; fIrsT fIrsT gONOcOXITEs ( FIg. 5 View Figure : fgc) largE, brOad, MEdIal MargIN sTraIghT, aNd pOsTErIOr MargINs caMbErEd; laTErOTErgITEs IX ( FIg. 5 View Figure : lT9) brOad, apIcally rOUNdEd aNd lONgEr ThaN apIcEs Of EIghTh laTErOTErgITEs aNd sTErNITE TEN. FUsEd valvIfErs IX ( FIg. 5 View Figure : vf9) brOadly EXpOsEd bETwEEN laTErOTErgITEs IX IN rEsT.

Measurements (in mm). Body length 13.1–14.0 (♀) / 11.5–12.5 (♂); length of head 1.95–2.4; lengths of aNTENNal sEgMENTs (I) 0.25–0.3: (II) 1.9–2.05: (III) 0.85–0.95: (IV) 1.35–1.45: (V) 1.2–1.35; MEdIaN lENgTh Of prONOTUM 2.5–2.75, hUMEral wIdTh bETwEEN apIcEs Of hUMEral prOjEcTIONs 6.55–6.70; lENgTh Of scUTEllUM 4.35– 4.45, wIdTh Of scUTEllUM 3.35–3.55.

Habitat. A. custos OccUrs  prIMarIly IN lOwlaNds, aNd has bEEN capTUrEd aT hIghEr alTITUdEs ONly spOradIcally. A ThErMOphIlOUs spEcIEs, IT Is MOsT cOMMON ON dry slOpEs, IN parks, aNd IN Orchards ( STEhlík 1987), bUT Is NOTAbout NOT rarE IN MOrE-hUMId placEs EIThEr ( SchUMachEr 1910). IT Is arbOrIcOlOUs, aNd EspEcIally cOMMON ON Alnus  spp. ( A. glutinosa  , A. incana  ) ( SchUMachEr 1910; HOffMaNN 1948; SEIdENsTückEr 1975; STEhlík 1987), bUT alsO frEqUENTly fOUNd ON Betula  aNd Tilia  spp. aNd lEss frEqUENTly ON Corylus  , Salix  , Populus  , Quercus  , Crataegus  , Betula  , Frangula  , Lonicera  , aNd Ailanthus  ( SchUMachEr 1910; SzUlczEwskI 1913; STEhlík 1987; STEhlík & VavříNOvá 1994), aNd alsO OccUrs ON cONIfErs ( STEhlík 1987; REUTEr 1909; SchUMachEr 1910; STEhlík & VavříNOvá 1994).

IN GErMaNy, IT OvErwINTErs IN ThE adUlT sTagE, aNd larvaE OccUr IN JUly aNd AUgUsT ( SchUMachEr 1910; WagNEr 1966). IT Is EspEcIally cOMMON ON Alnus  , whErE IT fEEds ON Agelastica alni  (L.) ( COlEOpTEra  : ChrysOMElIdaE  ) ( DUpUIs 1949; STEhlík 1987; LIs 2000) aNd ON Ulmus  ( AhMad & ÖNdEr 1990).

Molecular result. ThE UNUsUal MOrphOlOgIcal varIabIlITy IN hUMEral aNglE raIsEd ThE qUEsTION as TO whEThEr ThE TwO spEcIEs wErE IN facT ThE saME spEcIEs. AfTEr cOMparINg ThE gENITalIa Of ThE TwO spEcIEs, wE fOUNd ThaT ThEy wErE vEry sIMIlar IN MOsT rEspEcTs, sO MOlEcUlar daTa was NEEdEd. TwENTy COIAbout COI sEqUENcEs rEprEsENTINg fOUr spEcIEs wErE ObTaINEd. NO INsErTIONs Or dElETIONs wErE dETEcTEd, INdIcaTINg ThaT ThE sEqUENcEs wErE NOTAbout NOT psEUdOgENEs. All sEqUENcEs frOM OUr sTUdIEs wErE NEw aNd addEd TO ThE barcOdE daTabasE fOr ThIs grOUp, as wE cOUld NOTAbout NOT fINd cONspEcIfIc sEqUENcEs whEN a BLAST NUclEOTIdE sEarch was cONdUcTEd (GEN-BaNk accEssION NUMbErs arE lIsTEd IN TablE 1).

WIThIN ThE EXaMINEd COIAbout COI sEqUENcEs, a 574-bp rEgION was aNalyzEd; 445 sITEs wErE cONsErvEd, 129 sITEs wErE varIablE, 62 sITEs wErE parsIMONy-INfOrMaTIvE, aNd 67 sITEs wErE sINglETON sITEs. ThE aNalysIs Of NUclEOTIdE cOMpOsITION shOwEd ThaT ThE cONTENT Of (A+T) was 66.6%, whIch was hIghEr ThaN (G+C). ThIs rEsUlT was cONsIsTENT wITh ThE hIghEr (A+T) fEaTUrE Of ThE MITOchONdrIal gENOME NOTEd by HUa et al. (2008). ThIs aNalysIs shOwEd ThaT COIAbout COI had rEMarkablE cOdON UsagE bIas.

ThE INTErspEcIfIc aNd INTraspEcIfIc dIsTaNcEs basEd ON COIAbout COI wErE cOMpUTEd UsINg MEGA 7.0. ( TablE 2). ThE dIvErgENcE bETwEEN lONg-hUMErI ( A. custos  ) aNd shOrT-hUMErI ( A. chinensis  ) basEd ON COIAbout COI was 0.01247; hOwEvEr, ThE dIvErgENcE bETwEEN lONg-hUMErI ( A. custos  ) aNd A. koreana  , aNd ThaT bETwEEN shOrT-hUMErI ( A. chinensis  ) aNd A. koreana  wErE 0.10789 aNd 0.11130, rEspEcTIvEly. ThEsE wErE all apprOXIMaTEly TEN TIMEs hIghEr ThaN ThE dIvErgENcE bETwEEN lONg-hUMErI aNd shOrT-hUMErI. WhEN wE cOMpUTEd ThE dIvErgENcE Of ThrEE spEcIEs Of gENUs Arma  wITh P. lewisi  , ThE gENETIc dIsTaNcEs bETwEEN ThE TwO gENEra wErE IN ThE raNgE Of 0.17070–0.19725, dEpENdINg ON ThE Arma  spEcIEs UsEd. ThE ThrEshOld fOr a barcOdINg gap Is ThaT INTErspEcIfIc dIvErgENcE shOUld bE aT lEasT TEN TIMEs ThE avEragE Of ThE INTraspEcIfIc gENETIc dIsTaNcEs, accOrdINg TO HEbErT et al. (2004). ThErEfOrE, ThE gENETIc dIsTaNcE Of ThEsE TwO spEcIEs Is IN accOrdaNcE wITh ThE rEsUlTs Of MOrphOlOgIcal aNalysIs IN ThaT ThE dIffErENcE bETwEEN shOrT-hUMErI ( A. chinensis  ) aNd lONg-hUMErI ( A. custos  ) shOUld bE TakEN as aN INTraspEcIfIc varIaTION.

ThE aNalysIs Of NJ, BI, aNd ML TrEEs yIEldEd TwO MajOr cladEs wITh hIgh bOOTsTrap sUppOrT: cladE I Is rEprEsENTEd by TwO sEqUENcEs Of A. koreana  ( FIg. 6 View Figure ), cladE II INclUdEs 17 sEqUENcEs Of lONg-hUMErI ( A. custos  ) aNd shOrT-hUMErI ( A. chinensis  ), aNd Picromerus lewisi Is UsEd  as OUTgrOUp. ThE phylOgENETIc TrEEs prOvIdEd hErE arE ONly TO sEgrEgaTE TwO spEcIEs basEd ON ThE rEcIprOcal MONOphyly crITErIa, aNd arE NOTAbout NOT MEaNT TO dEfINE ThE phylOgENy Of ThE gENUs Arma  .


Zoological Museum, University of Copenhagen


Museum National d'Histoire Naturelle


Nottingham City Natural History Museum


Department of Biotechnology


University of Coimbra Botany Department














Arma custos ( Fabricius, 1794 )

Zhao, Qing, Wei, Jiufeng, Bu, Wenjun, Liu, Guoqing & Zhang, Hufang 2018


Cimex custos

Fabricius, 1794 : 94


Arma custos var. fuscoantennata

Motschulsky, 1860 : 501


Arma custos

Stichel 1961a : 656


Arma neocustos Ahmad & Önder, 1990 : 8

Ahmad & Önder, 1990 : 8 Thomas 1994 : 165


Arma neoinsperata Ahmad & Önder, 1990 : 9

Ahmad & Önder, 1990 : 9 Thomas 1994 : 165


Arma chinensis

Fallou, 1881 : 340


Arma discors

Jakovlev, 1902 : 64 Derzhansky et al. 2002 : 367 Kiritshenko 1922 : 14