Revisional notes on selectedselected EastEast PalaearcticPalaearctic species of AgabusAgabus Leach (Coleoptera, )Dytiscidae)

ANDERS N. NILSSONNILSSON

Nilsson, A.N.: RevisionalRevisional notesnotes onon selectedselected East PalaearcticPalaearctic speciesspecies of AgobusAgabus Leach (Coleo- ptera, Dytiscidae).Dytiscidae). [Taxonomiska[Thxonomiska anmiirkningaranmärkningar om östpalearktiskaostpalearktiska arter av Leach (Coleoptera, Dytiscidae).] -— Ent. Tidskr.Tidskr. I lll:149-161.1 1:149-161. Umeå,UmeA, SwedenSweden 1990.1990. ISSNISSN 0013-886x.001 3-886x.

Out fromfrom thethe studystudy ofof somesome EastEast PalaearcticPalaearctic speciesspecies of the ,4gabusAgabus Leach notes are pre- sented on classificationclassification and nomenclature.nomenclatwe. Agabus inexspectatusinexspectalus sp. n.n. isis describeddescribed from a single male from Siberia. The following syn. n. are presented: Agabus zaitzewi Poppius, 1909,1909, Agabus hudsonicushudsonlcus Leech, 1938, and Agabus mongolicusmongolicas Guorguiev,Gu6orguiev, 19681968 = Agabus pallens Poppius, 1905;1905; Eriglenus charini Lafer, 19881988: = Ilybius mandsuricus Guignot, 1956;1956; AgobusAgabus insignislnsignis Gu&nsuiev,Gu6orguiev, 1969 1969: = arcticusarcticus Paykull, 1798;1198; AgobusAgabus kurilensiskurilensis Kamiya, 19381938= = /rrslrs Aub&Aub6, 1838;1838; Mesogabus GtAorguiev,Gu6orguiev, 19691969= = GaurodytesGaurodyles Thomson, 1859.1859. Lectotypes are designated for thethe followingfollowing species:species: AgobusAgabus bergibergiZaitzev, Zaitzev, 1913,1913, Agabus tibetanustibetonus Zaitzev, 1908,1908, Agabus zimmermannizimmermonni Scholtz, 1920,1920, Colymbetes costulatus Mot-Mol- schulsky, 1859.1859. Agabus bergibergl isis given specificspecific rank.rank. Identification keyskeys are given for the world speciesspecies of the ,4A. . labiatuslabiatus (Brahm)(Brahm) group, andand forfor thethe malesmales of thethe PalaearcticPalaearctic speciesspecies of the .4.A. confinisconfi n is (Gyllenhal)(Gyllenhal) group.

A.N. Nilsson,Nitsson, DepartmentDepartment ofof AnimalAnimal Ecology,Ecology, UniversityUniversity of Umeå,Umed, S-901 8787 Umeå,Umed, .

Introduction The dytiscidd1'tiscid genusgems Agabus Leach, 1817,1817, includes is monophyletic. InIn current usageusage thisthis is most pro- almost 200 species, of which thethe majority are con- bably not thethe case,case, andand no no single single synapomorphysynapomorphy fined to thethe Holarctic Holarctic Region. Region. SharpSharp (1882)(1882) re-re- have been given for the largelarge setset ofof speciesspecies so clas- cognized 23 species-groupsspecies-groups in Agabus,Agobus, with some sified. groups "clearly artificial". LaterLater attemptsattempts ofof clas-clas- In thethe absenceabsence of thethe analysesanalyses asked for aboveabove sification have created a longlong listlist ofof subgeneric subgeneric a valuablevaluable task is to definedefine naturalnatural groupsgroups withinwithin names (Larson(Larson 1989:865).1989:865). However, the currentcurrent Agabus. A first stepstep inin thisthis directiondirection waswas recentlyrecently use of subgenerasubgenera shows a pronounced variation, taken byby NilssonNilsson && LarsonLarson (1990)(1990) in a revisionrevision of both personallypersonally and geographically. AfterAfter Sharp's thethe A. affinisaffinis (Paykull)(Paykull) group. AA classificationclassification of (1882)(1882) monography,monography, nono workwork has dealt with the the NearcticNearctic speciesspecies waswas recentlyrecently presentedpresented byby classification of Agabus out fromfrom thethe worldworld fau-fau- Larsson (1989),(1989), whose species-groupsspecies-groups names are na. However,However, ZimmermannZimmermann (1919)(1919) studied a large adopted here.here. TheThe delimitationdelimitation ofof groupsgroups callscalls for part ofof itit andand classifiedclassified the current subgenussubgenus Gau-Gaa- better knowledgeknowledge of many species,species, as many valu- rodytes Thomson, 1859,1859, into three groupsgroups fromfrom able characters were not givengiven withwith thethe originaloriginal structure ofof malemale genitalia.genitalia. descriptions or in handbooks. The prevailing regional perspective has probab- In thethe presentpresent paper,paper, somesome Agabus groups are ly hamperedhampered development,development, and anotheranother basicbasic discussed outout from the studystudy of EastEast PalaearcticPalaearctic problem is the position of AgobusAgabus as the dumping specimens and species.species. Some newnew synonymssynonyms are ground of Agabini.Agabini. ItIt seemsseems logicallogical that a genusgenus established and aa newnew speciesspecies is described from Si-Si- defined only on its wanting of the apomorphies of beria. MostMost of thethe materialmaterial forfor thisthis investigation investigation the other generagenera ofof thethe tribetribe shouldshould bebe impossibleimpossible was examined oror borrowed during a visit to the to classify. The first step towardstowards a natural classifi-classifi- Zoological Institute of LeningradLeningrad inin AprilApril 1988.1988. cationcalion of AgoDasAgabus must be to ensure that the genus The followingfollowing abbreviations are used:used: (MZH)(MZH)

150150 Anders N. NilssonNi/sson

University Zoological Museum, Helsinki, (ZIL)(ZIL) at leastleast on outerouter halfhalf ofof disc.disc. Metafemur Metafemur withwith produced. Zoological Institute, Leningrad; (TL) totaltoral length, posteroapical angle strongly produced. Elytron with punctures at intersections of meshesmeshes and in (MW)(MW) maximum width, (WC/WS) ratioratio betweenbetween longitudinallongitudinal series ofof about samesame size.size. MaleMale width of metacoxa and width of metasternum (cf. profemur without ventral brush of setaesetae ...... Nilsson 1984).1984). When possible, measurements are fulvaster 5. Metasternum with anteromedian impression given as mean -+± SD. 5 il;i;;i;;;;; ;l;il' ;;;;;;;;';;';;;;;*'#f"" long, attainingattaining levellevel of hind marginmargin of meso-meso- coxa. ProsternalProsternal processprocess narrower; ratio be-be- tween maximum andand minimum width 2.1-2.3.2.1-2.3 The Agabus labiatuslabiatus group tween Dorsal surface normally brown to piceous ...... お This group corresponds to the subgenussubgenus Eriglenus ¨…………………………・………… labiatus″b″ ′ — Metasternum with anteromediananteromedian impressionimpression Thomson, 1859.1859. It isis recognizedrecognized on thethe apicallyapically short, notnot attainingattaining levellevel of hindhind marginmargin of me- lobed parameresparameres and thethe narrow narrow metasternal metasternal socoxa.socoxa, Prosternal process broader; ratio be-be- wings, and includesincludes thethe sixsix species:species: A. fulvasterfulvaster tween maximum andand minimum width 2.4-2.9.2.4-2.9. Zaitzev, 1906,1906, A. labiatus labiatus (Brahm, 1790), 1790), A. Dorsal surface pale yellowish to reddish brown :::::i :::1i:: ::': rll::::l :: ll1llil litlt,,,,, pallens mandsuricus (Guignot, 1956),1956), A. pallens Poppius, 1905,1905, A. undulatusundulatus (Schrank, 1776),1776), andand A. zim-z,im- Agabus pallens Poppius mermanni Scholtz, 1920.1920. The species of the labia-labia- tus-group,r.s-group cancan bebe separatedseparated withwith thethe followingfollowing key.key. Agabus femoralrbfemoralis Payk. var.var. pallens Poppius, 1905:221905:22 Leech (1942) includedincluded alsoalso the Holarctic A. bi-bi- Agabus zaitzewi Poppius, 1909:12,1909 12, syn. n. Agabus hudsonicus Leech, 1938:123,1938:123, syn. n. farius (Kirby)(Kirby) in this group. However, as the para- farius Agabus mongolicus, Gui.orguiev,Gueorguiev, 1968:27,196827, syn. n. meres of thisthis speciesspecies are markedly different and the narrownarrow metasternal metasternal wings wings obviously obviously havehave TypeIlpe material. Holotype cta of pallens in MZH (Biström(Bistr6m evolved independently in severalseveral Agabus lineages, 1987),1987), labelled:labelled: "Olekminsk","Olekminsk", "Fl."Fl. Lena S.", "B. Pop- pius", "Mus. Zool.Zool. H:forsH:fors Spec. Spec. typ.typ. No No 1648 1648 AgabusAgabus Leech's classification is not followed here. Larson (Eriglenus)(Eriglenus) femoralisfemoralis Payk.Payk. v. pallenspallens Popp.",Popp.", and mymy (1989) (1989) placed A. bifariusbifarius in itsirs own group. holotype label. — LectotypeLectotype cra and paralectotypes 2Q29 of zaitzewizoitzewiinMZH in MZH- (Biström(Bistrom 1987)1987) labelled:labelled: "Kanin","Kanin", "B. Poppius", and mymy typetype labels;labels; lectotype lectotype alsoalso labelledlabelled Key to speciesspecies "zaitzewi"zaitzewi typ.typ. m.,m., A. salinatorsalinator typ. m.", and "Mus."Mus. Zool. H:forsH:fors Spec.Spec. typ.typ. NoNo 18511851 Agabus (Gaurod.) zait- 1.l. Elytron dark brownbrown toto blackblack withwith yellowyellow fasciafascia zewi Popp."Popp." Typesppes ofof hudsonicus andand mongolicus notnot or vitta,vitta, andand meshesmeshes ofof microreticulationmicroreticulation smallsmall seen in California Acad. Sci., SanSan Francisco,Francisco, and Mu- and roundedrounded ...... 22 seum of Nat.Nat. Hist.,Hist., Budapest, Budapest, respectively. respectively. - Elytron of moremore oror lessless uniform uniform yellowyellow toto brown colour, and meshesmeshes of microreticulationmicroreticulation TypeIlpe localities: of pallens "Olekminsk""Olekminsk" (USSR,(USSR, Yakuti-Yakuti- of more irregular sizesize and shape 33 ...... an ASSR);ASSR); of zaitzewi "Tschosha", KaninKanin PeninsulaPeninsula 2. Elytron Elytron with with yellow yellow marginalmarginal vitta that inin poste- (USSR,(USSR, Nenets National Okrug); of hudsonicus "Chur-"Chur- riorrior 2/3 isis split by narrow black stripe. Anterior chill, Manitoba" (Canada);(Canada); of mongolicus "Chentej"Chentej ai- margin of metacoxametacoxa sublaterallysublaterally not fully re-re- of not fully mak, 2020 kmkm SW SW von von Somon Somon Norovlin" Norovlin" (Mongolian (Mongolian aching level ofof posterior margin of mesocoxa of ... People's Republic).Republic). mandsuricus — Elytron normally with;iii; subbasal and submedian ;i;,;;; ;;;ii; ;;;;;i ;;; ;,ff;xf'#"" Note. The holotype of pallens was described as a male. yellow vittae andand subapical spot. Anterior mar- Anterior mar- However, as the specimenspecimen referred to aboveabove otherwiseotherwise gin ofof metacoxametacoxa sublaterallysublaterally reachingreaching slightlyslightly fits the description itit was probably onlyonly erroneously sex-sex- anterior to levellevel of posteriorposterior marginmargin ofof meso-meso- ed by Poppius. InIn 19831983 II provided thethe typetype seriesseries of zait- coxa ...... ,undulalusundulatus zajt- zewi withwilh my lectotype and paralectotype labels. How- 3. Body Body elongate elongate ovaloval withwith totaltotal lengthlength 7.07.0 mm or ever, by mistake these specimens were published as holo- less. Ventral face of metatibia micropunc- less. Ventral face of metatibia either micropunc- type andand paratypes paratypes respectively respectively (Larsson(Larsson && NilssonNilsson tate or with macropunctures alsoalso on disc ...... 44 . 1985:126).1985:126). InsteadInstead my lectotype and paralectotype de- — Body broadly oval with total length Body broadly oval with total length about 8.0 signations werewere publishedpublished byby BiströmBistrom (1987:38).(1987:38). mm. VentralVentral face of metatibiametatibia withoutwithout micro-micro- punctation and disc without punctation and disc without macropunctures ... Larson & NilssonNilsson (1985,(1985, as A. zaitzewi) recog-recog- zimmermanni zoitzewi) 4. Ventral v;;r;;i' facei;;; of;i metatibia;ili;i; without;; :;;; macropunctu-;;,?#tr::ff: "' nized the Holarctic distribution ofof thisthis speciesspecies and res on disc. Metafemur with posteroapical angle also noted thatthat itit eventually eventually waswas conspecificconspecific with weakly produced. Elytron with punctures at in- A. labiatus,lobiatus, as also judged byby PoppiusPoppius (1905)(1905) who tersections of meshesmeshes much smaller than serialserial described it as a colour form of ,4A. . labiatus.lobiatus. Later, punctures. Male profemur with ventral brush of (1943) setaesetae...... ,...... ...... 55 Brinck (1943) reportedreported on specimensspecimens from Mon- — Ventral face of metatibia with macropunctures golia and Siberia olof the same pale colour and esta- Notes on Eost Palaearctic Agabus l5l ― blished the subspecies A. lobiotus pallens Pop- pius. Besides the testaceous dorsum, Brinck (1943) ― ― noted upon the larger size and the larger meshes of 洲 the more deeply impressed elytral reticulation of this subspecies. 鮒 The study of the holotype of A. pallens has con- ― ― ― ― ２ firmed the identity of this species with ,4. zoitzewi, and not with .4. labiotus as previously accepted. ― ― ― ‐ The most important characters for the separation of this species-pair are: (l) the development of the anteromedian emargination on the metasternum - deeper in A. labiotus; and (2) the width of the prosternal process - broader in A. pollens. ln contrast to A. lobiatus, the females of A. pallens frequently have the elytral reticulation very deeply impressed with punctures at intersections reduced. Figs l-3. Agabus zimmermanni Scholz. Male genitalia. There is some variation in the shape of the median Scale bar 0.5 mm. - I . Paramere, external aspect. -2-3. lobe of the aedeagus, but the few males of A. pal- Median lobe of aedeagus. -2. Dorsal aspect of distal 2/1. Right lateral aspect. /ens available do not permit a separation of indivi- -3. dual and specific differences. Gu6orguiev (1968) assigned his ,4. mongolicus men of A. zimmermonni was found misidentified to subgenus Gaurodytes, species-group A. nebu- as A. labiatus. The specimen is labelled "Brato- /osrus (Forster). However, my study of the female lyubov. Aleksandr. u. (A. Jacobson)" (here tran- specimens from Mongolia reported on by Gudor- scribed). According to Dr. I.M. Kerzhner, Lenin- guiev (1969) and additional material in ZIL have grad, the locality is Bratolybovka, former in Alek- shown that A. mongolicus is a true member of the sandriyskiy uezd, now in Dolinskiy rayon of Kiro- labiatus-group and identical with 1. pallens. vogradskaya oblast in Ukrainskaya SSR (appr. 48' Additional specimens studied of A. pallenswere l5'N 32' 45'E). If the specimen was correctly collected by Dr R. Angus in Siberia: by Lena river labelled, this is the first European record of .4. to 140 km upstream from Yakutsk 27-29.vi.1970, zimmermanni, and also the first record from the 4?; at Tibelti in Irkut valley 28 km W of Lake Bai- USSR. Another interesting record of this species is kal 8-15.vi.1970, l9; Viluiskiy Tiakt to 55 km W from the Black Sea coast of , Nessebar of Yakutsk, 2-5.vii.1970, lcr39. A male of A. la- 26.vi.1965, leg. T. Palm. biotus $tas taken by Dr Angus at Karasuk in No- vosibirsk oblast 28.v.-28.vi.1982 together with ,4. Agabus mandsuricus (Guignot) fulvaster. Evidently, A. pallens has a wide distri- bution in the Palaearctic Region from the Kanin Ilybius mandsurrcus Cuignot, 1956a:139; 1956b:396, 397. Peninsula to East Siberia and Mongolia. Eriglenus chorini Lafer, 1988:54-56, fig. 2.

(1986) Agobus zimmermanni Scholz Wewalka transferred this species from lly- biusto Agabus, subgenus Eriglenus, and provided Figs l-3. a detailed description with illustrations of male genitalia. Lafer (1988, 1989) was obviously not Agabus zimmermanni Scholz, 1920:15, 16. aware of Guignot's species when he described E. Type material. Lectotype cr here designated, labelled: charini. I have not seen Lafer's types, but the ori- 'Asia Minor", "cr", and my lectotype label. In coll. ginal description with illustrations leaves no doubt Zimmermann, National Zoological Collection, Munich. on the identity of this species. This rare species was previously known only from Asia Minor (Zimmermann & Gschwendtner 1935: The Agabus confinis group 13, Zaitzev 1953:269\. Besides the types series I have seen a specimen collected by Dr G. Wewalka This group has earlier been referred to as the /. in Turkey, prov. Aydin 27 Jv.1969. ln ZIL a speci- congener (Thunberg) group (Zimmermann 1934, 152 Anders N. NrTsson

Nilsson 1984). Larson's (1989) renaming of the Aedeagus with subapical spine pointed ...... 2 with ventral tooth. Antenna group is accepted as it is based on a well-defined 2. Protarsomere 5 subserrate. Prosternal process flat. Aedeagus as Holarctic species. The confinis-group in my sense in Fig. 8 ...A. elongatus (Gyllenhal) includes also the elongqtus- and zetlersledti- Protarsomere 5 withouth ventral tooth. Anten- groups of Larson (1989). In the Palaearctic, the na simple in most species. Prosternal process in most species ofat least low convexity ...... 3 co nfi n is-, j apon icus- and fusc ipe n nis-groups are 3. Elytron with a metallic sheen; lines of reticula- all characterized by the bifid median lobe of the tion coarse and meshes longitudinally stretched aedeagus. These groups also share the following basally. Aedeagus as in Fig. I I ...... characters: (l) disc of pronotum black and con- trasting with variously paler elytra, (2) clypeus E i ;i;; ;'; i; h; ; ; ;;;i ii;,f;.'lf Xl:J,I.(,J ; i?!l i-"" culation not longitudinally stretched ...... 4 with anterior bead more or less continuous, (3) 4. Elytron with strong subsutural row of punc- metasternum with anteromedian emargination tures that almost reaches base. Epipleuron dark long and attaining level of hind margin oi meso- piceous to black, at least in inner third. Aede- coxa, (4) elytron with simple reticulation, (5) me- agus as in Fig. l0 ...... A. tibetonus Zaitzev Elytron with subsutural row of punctures weak. posteroventral tatibia with margin impunctate or Epipleuron testaceous to rufopiceous ...... 5 with few well-separated punctures, (6) pronotum 5. Metasternum and -coxae medially densely with anterior bead broadly interrupted medially, punctate and with long conspicuous setation. and (7) paramere stylate. Within the aggregate, Aedeagus strongly curved in lateral view (Fig. 12) ...... A. setulosus (J. Sahlberg) the fuscipennrs-group is characterized by the Metasternum and -coxae medially with at most broadly dilated male protarsus, with tarsomere I occasional punctures and without setation. twice or more as wide as tarsomere 5. The japoni- Aedeagus less strongly curved in lateral view .... 6 cr,,s-group is characterized by the short and robust 6. Aedeagus with subapical spine very thin and not protruding (Fig. 20) hind legs with tarsomere 5 only about longer l/3 A. zet terstedt i Thomson than tarsomere 4, and pronotum with hind angles Aedeagus with subapical spine broader and posteriorly produced and slightly acute. I have protruding ...... 7 found ny synapomorphies for the confinis-group 7. Aedeagus not broadened at level of straight subapical spine (Figs l7-19) ...... 8 of species, in which distinct evolutionary novelties Aedeagus more or less broadened at level of are restricted to single species. The problem of straight or curved subapical spine...... l0 how to deal with this group demands further stu- 8. Apical segment of maxillary palpus infuscate. dy. The separation of it into smaller units will still Body more elongated with sides subparallel. Pronotum not so broad posteriorly, with great- with end up a large rest-group, and the alternative est width sometimes anterior to hind angles and to fuse the three groups discussed above should al- width at hind angles/width at front angles so be considered. In this case the subgeneric name about 1.4. Aedeagus as in Fig. 17 ...... Acotodes Thomson, 1859, is available. A. moestus (Curtis) Apical segment of maxillary palpus at least ba- here present genitalia I will drawings of male sally testaceous. Body with lateral outline more (Figs 4-20) and give a key to the Palearctic species curved. Pronotum broader posteriorly, width at of the confinr.s-group. After the key some notes hind angles/width a front aaglesabout 1.6...... 9 on and nomenclature are given for a 9. Body length 8.3-9.5 mm. Metasternal wing broad, WC/WS 1.9-2.2. Aedeagus as in Fig. l9 few poorly species. known A new species is de- . A. confinis (Gyllenhal) scribed lrom Siberia that is provisionally assigned Body length 6.7-7.6 mm. Metasternal wing to this group as it differs from all other species in narrow, WC/WS 2.4-2.8. Aedeagus as in Fig. the shape of the aedeagus that lacks a ventral sub- 18...... A. levanderiHell€n Protarsal claws markedly shorter than protar- apical spine. I have not seen .4. turcmenus Guig- somere 5. Prosternal process very slightly con- not, 1957, that probably belongs to the confinis- vex, almost flat. Metasternal wing narrow, group. It was described from Tirrkestan and dif- WC/WS 2.7-2.9. Body length 6.0-7.0 mm, la- fers from most other species in its large size, TL teral outline strongly curved ...... I I 一 Protarsal claws subequal to or slightly longer 9.5-10.0 mm (Guignot 1957). Ｈ than protarsomere 5. Other characters varying . l2 Elytron with meshes of reticulation small, Key to males of Palaearctic species rounded and regular in size. Aedeagus as in Fig. l5 ...... A. clypealis (Thomson) l. Aedeagus with subapical spine broadly rounded Elytron with meshes of reticulation larger and (Fig. 9) ...... more irregular in size and shape. Aedeagus as in . sp. n. from Hokkaido and Sachalin (Sat6 in prep.) Fig. I 6 ...... A. pseudoclypeo I is Scholz Notes on EastEast PalaearcticPaloeorclic AgabusAgobus 153153

12.12. AedeagusAedeagus stronglystrongly broadenedbroadened atat levellevel of broad, not aa subspeciessubspecies ofof,4. A. congener.congener. The diagnosticdiagnostic ventrally curved subapicalsubapical spinespine (Figs(Figs 4, 13)l3) ..... 13l3 characters areare listedlisted below.below. — Aedeagus sligthly broadened at levellevel of usuallyusually Male genitalia:genitalio: narrower subapical spine (Figs 5-7)5-7) ...... 14l4 Male aedeagus (Fig. 13)13) shorter andand 13.13. ProsternalProsternal processprocess ofof low low convexity, convexity, almostalmost more strongly curved in lateral viewview than inin A.,4. flat. BodyBody elongate,elongate, TL/MWTL,/MW 1.9-2.0.1.9-2.0. Aede-Aede- congener, subapical spine very broad; paramereparamere agus shorter andand moremore stronglystrongly curvedcurved (Fig.(Fig. 13)l3) (Fig.(Fie. 14)la) rather broad. .... A. bergibergi Zaitzev Body size and shape:shape: the six specimensspecimens measured — Prosternal process distinctly convex in crosscross Body size ond the six section.section. BodyBody broader,broader, TL/MWTL,/MW 1.7-1.8. 1.7-1.8. are of aboutabout thethe samesame sizesize as A. congenercongener (TL(TL 7.1 Aedeagus longer and lessless strongly curved (Fig. ±-r 0.30.3 mm),mm), butbut thethe shapeshape isis muchmuch narrowernarrower andand 4) ...... A. congenercongener(Thunberg) (Thunberg) thus similarsimilar to A. thomsoni thomsoni (TL/MW 1.96 ±-t 14.14. BodyBody lengthlength 7.9-8.5 mm.mm. TL/MWTLIMW 1.8-1.9.1.8-1.9. 0.02). Aedeagus as in Fig. 7 ...... ...... A. costulatuscoslulotus (Motschulsky)(Motschulsky) Colour:Colour: pronotum blackblack withwith onlyonly narrow laterallateral - Body length normally lessless than 7.87.8 mm,mm, bead rufous; elytron rufous. TL/MWTLIMW eithereither slightlyslightly largerlarger or smallersmaller ...... 15l5 OtherOther characters:characters: prosternal process of lowlow con-con- 15.15. BodyBody lengthlength inin most most specimens specimens larger,larger, TLTL vexity, similar to A. clypealis; metasternal wing of 6.8-7.8 mm.mm. Body Body moremore elongated,elongated, TL/MWTLlMw vexity, similar to A. clypealis; metasternal wing of 1.9-2.0.1.9-2.0. AedeagusAedeagus asas inin Fig.Fig.6 6 ...... moderate width, WC/WS 2.51 ±-f 0.19;0.19; elytronelytron A. thomsonithomsoni (J.(J. Sahlberg)Sahlberg) with meshes of reticulation irregular and relativelyrelatively - Body length in mostmost specimens specimens smaller,smaller, TLTL wide. 6.3-6.9 mmmm BodyBody broader,broader, TLIMWTL/MW 1.7-1.8.1.7-1.8. Aedeagus as in Fig. 55 .....A.A. lapponicus/appon rcus (Thomson)(Thomson) Distribution. A. bergibergi is so farfar knownknown onlyonly fromfrom the highlands of TranscaucasiaTianscaucasia in the Gruzinskaja Agabus bergibergi ZaitzevZaitzev and ArmjanskajaArmjanskaja SSR. SSR. Some Some localities localities werewere givengiven by Zaitzev (1927:26). The presence of A. congenercongener Figs 13, 14.14. in thisthis regionregion is uncertain. TheThe singlesingle femalefemale stud- Agabus bergibergi Zaitzev, 1913:195,l9l3:195, 196.196. ied (from(from Baraleti)Baraleti) ofof whatwhat Zaitzew Zaitzew (1927)(1927) listedlisted GaurodytesCaurodytes congenercongener bergi Zaitzev, 1953:248.1953:248. as A. congenercongener is probably A. pseudoclypealis.pseudoclypealis. TypeTtpe material (in ZIL).ZIL). Lectotype ct (here(here designated;designated; genitalia extruded) labelled "Lac. LevangölLevangtit 8500'8500'prov. prov. AgabusAgobus costulatuscostulotus (Motschulsky) Tifl. 25.VI.09 Schmidt" and mymy lectotypelectotype label. — Pa- ralectotypes 2Q,29, oneone withwith samesame originaloriginal labellabel asas lectoty-lectoty-- Fig.Fig.7. 7. pe and the other one labelledtabelled "Gr. oz.oz. Levan-gelLevan-gel Azjalg. ol. u.u. Tifl.Tifl. L. L. Berg Berg 25.VI.09" 25.v1.09" (here(here transcribed)transcribed) (head(head ColymbetesColymbetes costulatuscoslulatus Motschulsky, 1853:81853:8 (nomen nu- and prothoraxprothorax lost),lost), andand withwith mymy paralectotypeparalectotype labels.labels. dum). 1YpeIlpe locality:locality: "Transcaucasia:"Tianscaucasia: Prov. Tiflis, lacuslacus Le-Le- ColymbetesColymbetes obscuricollis Motschulsky, 1853:81853:8 (nomen vangöl,vangdl, 8.500' " (Grusinskaja(Crusinskaja SSR).SSR). nudum). ColymbetesColymbetes costulatus Motschulsky, 1859:541,1859:541, 542.542. Note. The original description was based onon lcr2Q1029 coll-coll- GaurodytesCaurodytes tunkunensislunkunensis Zimmermann, 1928:178,1928:178, 179.l'79. ected by BergBerg and SchmidtSchmidt and placedplaced inin thethe museummuseum collections in Leningrad andand Tbilisi.Tbilisi. EvidentlyEvidently ZaitzevZaitzev TypeIlpe material (in(in ZIL). Lectotype Q9 of ColymbetesColymbetes cos-cos' laterlater receivedreceived additionaladditional specimensspecimens collectedcollected togethertogether tulatustulatus (here(herc designated) labelled:labelled: "Jakutsk"Jakutsk 3156-1.", with the typetype seriesseries asas hishis collectioncollection inin LeningradLeningrad in-in- "3156"3156 costulatus Motsch." (in(in thethe handwritinghandwriting of Mot- cludes four specimensspecimens so labelled. I have selectedselected 1(729lcr29 schulsky)schulsky) and my lectotype and identificationidentification labels.labels. —- of thisthis seriesseries as types.types. Paralectotype 9 (here(here designated)designated) labelledlabelled "Jakutsk"Jakutsk 3156-2",3156-2", andand mymy paralectotypeparalectotype andand identificationidentification la-la- Zaitzev (1913) first placed A.,4. bergibergi nearnear A. lappo-lappo- bels. lype'Ilpe locality:locality: "Gouvernement"Couvernement de Iakoutsk"lakoutsk" (Jakutska-(Jakutska- nicus, from which it differed in the lighterlighter elytra, nicus, frorn it differed ja ASSR).ASSR). the more elongated body shape, and thethe broader,broader, Note. ZaitzevZaitzev (1905) reported on the presencepresence of syn-syn- less convexconvex prosternalprosternal process.process. LaterLater (Zaitzev (Zailzev typestypes of A..4. costulatuscostulolusin in the Zoological Museum, Lenin- 1953)1953) he treated A. bergibergi asas a subspeciessubspecies of A./4. grad. LaterLater (Zaitzev(Zaitzev 1910:37)l9l0:37) he stated that itit waswas 1(719lcrlQ from Jakutsk.Jakutsk. However,However, asas thethe paralectotype paralectotype female female congener (Thunberg),(Thunberg), confined to thethe Transcauca-Tianscauca- congener to lacks both thethe forefore andand midmid legs,legs, ZaitzevZaitzev probablyprobably mis-mis- sian highlands,highlands, and characterizedcharacterized by thethe above-above- identified it as a male. mentioned characters. To settle the status of A. bergi,bergi, the specimensspecimens Zaitzev (1905, 1910,1910, 1953)1953) listed the characterscharacters present in thethe ZaitzevZaitzev collection were studied. MyMy that separatedseparated A. costulatus costulatus fromfrom A. congener.congener. conclusion is that A.1. bergibergiis is a validvalid species,species, and However, in his 19531953 keykey A. costulatuscostulalas is placed 154154 ИAnders″αθrs N.Nl N7ノNilssonssO″

Figs 4-ll.4-11. Agabus, median lobe of aedeagus,aedeagus, rightright laterallateral aspect.aspect. ScaleScale barbar 0.50.5 mm. mm. —4. -4. A. congenercongener (Thunberg).(Thunberg). —5.-5.A. A. lapponicuslapponicus(Thomson)._-6.A.thomsoni(J.Sahlberg).-T.A.costulotus(Motschulsky). (Thomson). —6. A. thomsoni (J. Sahlberg). —7. A. costulatus (Motschulsky). —8.-B.A.elongatus A. elongatus (Gyllenhal).(Gyllenhal).-9.SpeciesfromJapantobedescribedbyProf.Sat6.-10. —9. Species from Japan to be described by Prof. Satö. —10. A.tibetonusZaitzev.--ll.,q.slovzolvi(J.A. tibetanus Zaitzev. —11. A. slovzovi (J. Sahlberg). Notes on East Polaearctic Agabus 155

Figs l2-20. Agabus, median lobe of aedeagus (12, 13, l5-20) and paramere (14), right lateral aspect. Scale bar 0.5 mm. -12. A. setulosus(J. Sahlberg). -13-14. A. bergiZaitzev.-15. A. clypealis(Thomson). -16. A. pseudoclypea- /is Scholz. -17. A. moestus (Curtis). -18. A. levonderi Hellen. -19. A. confinb (Cyllenhal). -20. A. zetterstedti Thomson. 156 ノ4′ グ ヾLノV′ /ssO′ `rs′ 1.9

３ thomsoni Σ ヽ ニ １ ８ ． ー costulatus ヨ lapponicus ト ¨ Ш ■ く 工

∽ １ ７ ＞ 口 Ｏ ｍ

16 60 6.5 7.0 75 8.0 BODY LENGttH,TL― h(mm) Fig. 21. Relationship between body length (TL-h) and body shape (TL-h/MW) in samples of ,Agobus costulatus (Motschulsky), A. lapponicus (Thomson) and A. thomsoni (J. Sahlberg). Data on A. lapponicus and A. thomsoni from Nilsson (1985). For A. costulatus mean t SD are given for ten Siberian specimens.

F6rhAllande mellan kroppsliingd (utan huvud) och kroppsform (liingd utan huvud/stOrsta bredd) fdr tre olika arter. Viirden ges som medelviirde + standardavvikelse. nearest to A. thomsoni, and as also noted by Gu6- Other characters: prosternal process of moderate, orguiev (1972) this is the species that is most diffi- even convexity; metasternal wing broadly triangu- cult to separate A. costulatus from. In this spe- lar, WC/WS 1.85 + 0.12, N:9; male protarsal cies-pair together with .4. lopponicus the male claws very slightly shorter than protarsomere 5; genitalia are more or less identical (Figs 5-7) and elytron with lines of reticulation fine, often coarse identification is so far only possible from biomet- in female, and meshes irregular in shape and rela- ric characters like body size and shape (Fig. 2l) tively large. (Nilsson 1985). Zaitzev's (1953:220-221) key cha- racters based on the size and number of serial A. costulatus has a wide distribution in the NE punctures on the elytron are not supported from Palaearctic (Zaitzev 1953, Gueorguiev 1972). It the material I have studied. This goes also for the should be sympatric with,4. thomsoni in the E Si- four characters, inclusive of aedegal shape listed berian tundra where the latter species has a rare by Gu6orguiev (1972). Some diagnostic characters occurrence according to Zaitzev (1953). The study are given below. of long series of local populations from this region could improve the current understanding of this Mole genitalia: aedeagus long and only slightly species-pair. broadened at level of narrow subapical spine (Fig. 7). Agabus tibetanus Zoitzev -t-0.3 Body size and shape: TL 8.2 mm, and Fig. 10. TLIMW 1.90 + 0.04, N:10. Colour: pronotum black with only narrow lateral Agabus tibetanus Zaitzev, 1908:425, 426. bead rufous; elytron rufotestaceous to rufopiceo- Ilpe material (in ZIL). Lectotype cr (here designated) la- us. belled "V. tja. Toso-nora 13.300! II.95 Rob. i Kozl." Notes on East Paloearctic Agobus 157

(here transcribed) and my lectotype label. Paralecto- types 6o5Q (here designated) with same original- labels and my paralectotype labels. Ilpe locality: "Tibet or: lacus Toso-noor, litus orient., 13.300' " (China, prov. Tsinghai).

This rare species is known only from the type se- ries. Within the confinis-group it is recognized on the following diagnostic characters.

Male genitolia: aedeagus (Fig. l0) short and ro- bust; only slightly broadened at level of narrow subapical spine. Body size and shope: relatively small, TL 6.6 -r 0.3 趙 mm; body only slightly narrower than in A. con- gener,TL/MW 1.83 i 0.03, N:ll. Colour: pronotum black with only narrow lateral bead rufous; elytron rufous; epipleuron dark pice- ous to black, at least in inner third. Other choroclers: prosternal process long with Figs. 22-24. Agabus inexspectatus sp. n., male genita- apex acute, more or less tectiform in cross-section; lia. Scale bar 0.5 mm. -22-23. Median lobe of aedea- metasternal wing broadly triangular, WC/WS gus. -22. Right lateral aspect. -23. Dorsal aspect of dis- tal 2/3. -24. Paramere, external aspect. 2.08 + 0.09, N: 10; elytron with meshes of reticu- iation large and irregular, and longitudinal series punctures of strong with subsutural row distinct latively narrow, anterior bead only present lateral- male protarsal almost to base; claws slightly long- ly; reticulation on disc with small meshes irregular protarsomere er than 5. in size and shape; elytron relatively shiny, longitu- dinal series of punctures distinct, reticulation with Agabus inexspectotus sp. n. meshes generally small and rather deeply engrav- ed, irregular in shape and size, and frequently Figs 22-24. with rather strong punctures at intersections; prosternal process Description of male glabrous; metasternal plate with rather fine irregular reticulation, meshes Body size ond shope; TL 6.0 mm, MW 3.2 mm; longitudinally stretched towards posterior margin; body elongate with lateral outline slightly curved, last abdominal sternum subrugose in posterior subparallel, TLIMW 1.85. half; metafemur ventrally with fine longitudinal Colour: head black with narrow anterior margin reticulation and spread micropuntures; metatibia and two posteromedian spots rufous; antenna and ventrally with fine transverse reticulation, disc im- palpi rufous; pronotum black with narrow lateral punctate, anteroventral margin with row of seven bead rufopiceous; elytron rufopiceous; ventral separated punctures in proximal 3,/4. surface mainly black with epipleuron, metacoxal Structurol character$ head broad, basal width processes and apical margin of abdomen rufous; about 2/3 of width of pronotum at base with sides abdominal sterna 2-4 medially rufopiceous; legs very weakly curved; prosternal process narrow rufous with femora rufopiceous. and tectiform, lateral bead well-defined; metaster- Sculpture and setation: clypeus with anterior bead num with anteromedian emargination attaining le- almost continuous, broken medially into closely vel of hind margin of mesocoxa; metasternal wing spaced transverse punctures; dorsal reticulation narrow, WC/WS 2.78:' pro- and mesotarsomeres simple, on head with small irregular meshes that 1-3 weakly dilated and ventrally with transverse occasionally have a central micropuncture, me- rows of small, rounded scales; protarsal claws shes wider and less deeply engraved on clypeus; elongated, slightly longer than protarsomere 5; pronotum with anterior submarginal row of punc- metafemur slender, posteroapical angle not pro- tures medially broken, gap subequal to length of duced posteriorly; metatarsomere I much longer antennomere 1l; lateral bead well-defined and re- than longer tibial spur. 158 Anders N. Ni/sson

Genitolia: aedeagus simple (Figs 22, 23), curved in notaticollis (Aub6, 1837:107). However, J. Sahl- lateral view, submedially broadened and subapi- berg (1880:55) interpreted Motschulsky's descrip- cally strongly narrowed, distally with ventral tion as referring to a species of Acatodes (now sy- ridge; paramere (Fig. 24) simple with inner and nonym of Agabus), which he described under the apical setation long. binomen Acatodes nigriventris, attributed to Mot- Female and biology unknown. schulsky. Zaitzev (1905) stated that A. nigriventris is a senior synonym of Agobus coxalls Sharp, Ilpe material. Holotype o in ZlL,labelled "Olekminsk 1882, and thus the valid name of the species. ,4. Jakutsk. ob. Gerts 30.V.901" (here transcribed). nigriventris was described from specimens in the Ilpe locality: Oljokminsk, Jakutskaja ASSR, Siberia. collection of Obert taken at Krasnojarsk by Streb- Derivation of name. The specific ephitet aims at the unexpected combination of characters found in this spe- lov. This collection is partly lost, and no such spe- cies. cimens could be found in ZIL. However, as Zait- zev (1905) stated that Sahlberg had compared spe- This small species has the habitus of the confinis- cimens of A. coxalis sent to him with the typical group, but the simple aedeagus contradicts group specimen of his nigrivenlrrs, at least one specimen membership. The aedeagus of A. inexpectatushas was kept by Sahlberg in his own collection. I have almost the characteristic apex of the group, and seen a female of A. coxalis from coll. Sahlberg in the subapical spine could easily have evolved from Zoological Museum, University of Turku, labelled a split of the ventral ridge. The shape of body and as nigriventris by Sahlberg. Another specimen in pronotum is reminiscent of A. elongalas, in my the Sahlberg collection, from E Siberia, has been opinion a species that belongs to a lineage that se- identified by Zaitzev as "Acothodes nigriventris parated from the rest ofthe group near its base. In Motsch.". Strangely enough, this specimen be- this case, the similarity with ,4. elongatus could in- longs to Agobus trrslrs Aub6. dicate that the new species evolved near the base It seems correct that Sahlberg's record of nigri- of the sister-group of the confinis-group. The re- ventris represents /. coxalis. However, Acatodes cognition of this sister-group is problematic as ,4. nigriventris is only a misidentification and conse- inexspectotus has no good apomorphic character quently the name cannot be used as a valid species relativ to the large group of Agabus species with a name (cl. Article 49 of the Code). continuous clypeal bead and a long metasternal A. coxalis is in the group recognized on the re- impression. The medial gap in the anterior row of duced subapical spine of the aedeagus. punctures on pronotum indicates a relationship with the A. guttotus (Paykull) group, but this is contradicted by the broad and flat prosternal pro- The Agabus arcticus group cess of this group. Zimmermann (1934) named this group after A. sturmii (Gyllenhal), but Larson (1989) changed the name to the arcticus-group in order to name The group groups after Holarctic species when possible. Lar- This is a small group with four species (Larson son (1989) restricted the group to three species, 1989), of which three are Holarctic. Only one of and discussed the position of A. zetterstedti that them is treated here. was placed in its own group (see above). Seeming- ly, a few more Palaearctic species like,4. conspi- caas Sharp and A. browni Kamiya show affinities Agabus coxalis Sharp to this group. Elytral sculpture and the aciculate sculpture of the metafemur unite this group with Acatodes nigriventris (Motschulsky) sensu J. Sahlberg, the /r,s/,s- and ambiguus-groups (Nearctic species 1880:55 (misident.). Agabus coxalrs Sharp, 1882:535. revised by Larson 1989), and the classification of Agabus splichali Reitter, 1899: 196. this aggregate should be studied further. A gabus sc hm idti Zailzev, 19 13 : 197 . Gourodytes ermaki Zaitzev, 1953l.259. Aga bus a rct icus ( Pay ku I l)

As noted by Zaitzev (1905), Rhantus nigriventris Dytiscus arcticus Paykull, 1798:201. Motschulsky, 1860:101, is a junior synonym of R. Colymbetes reticulolus Kirby, 1837:71. Notes on East Palaearctic Agabus 159

Colymbetes glocialis Motschulsky, 1853:7 (nomen nu- /us Guignot, 1954, A. debilipes Rdgimbart, 1899, dum). and A. solskii Jakovlev, 1897, belong to this Colymbetes alpinus Motschulslcy, 1860: 102. group. A ga bus subfasciatus LeConte, I 863 : I 7 (nomen nudum). Gaurodytes punctipennis J. Sahlberg, 1880:345. Gaurodytes sibiricus J. Sahlberg, 1880:345. Agabus ochoticus Poppius, 1908:54. Agabus tristis Aubd Agabus arcticus ab. collaris Scholz, 1917:251 (rank in- Agabus lnsrrs Aub6, 1838:356. frasubspeci fic). Agabus kurilensis Kamiya, 1938a:35, syn. n. Agobus insrgnrs Gu6orguiev, 1969:62-64 + figs l-3, syn, n. Larson (1989) listed more synonyms of this spe- When visiting ZIL I was surprised to find in the cies. However, Agabus subopacus Mannerheim collections some specimens of the well-known should be omitted from this list as it is a junior sy- Holarctic species A. arcticus from Mongolia iden- nonym of Rhontus suturellus (Harris) (Zaitzev tified by Gu6orguiev as A. insignis. A study of the 1906). original description of A. insignis made the syno- The identity of A. kurilensis has long been in nymy given above very likely. The study of the ho- doubt, and Zaitzev (1953:246) suggested that it lotype and two paratypes from the Zoological In- was only a colour variant of A. conspicuus. On stitute, Warszaw, has confirmed the synonymy. the contrary, Nakane (1989) listed A. kurilensis as Within the genus, this species is easily identified a valid species. Kamiya's collection, kept at the on the coarse, simple elytral sculpture, the flatten- Entomological Laboratory of the Tokyo Universi- ed prosternal process, and the denticulate anterior ty of Agriculture, was destroyed by fire during protarsal claw and the large round scales on the World War Two (Nakane, in lilt.). However, KOno ventral surface of protarsomeres 1-3 of the male. (1944) reported on additional specimens from the The colour of the dorsal surface varies from pale Kuriles collected in 1941. Two females from this yellow to black. In the former case the pronotum series were borrowed from the Entomological In- has at least a brown anterior margin and medioba- stitute of the Hokkaido University. The following sal spot, and in the latter at least the lateral prono- labels were attached to these specimens: "Kurilen tal margins are rufous. This colour variation is H. K6no S. Sumimiya 20.VII", "Agabus (Gauro- seemingly dependent on the habitat and shows no dytes) kurilensis Kamiya det H. KOno". Both fe- geographic pattern (cf. Larson 1989). The,4. in- males are identical with ,4. /rislls, as also made slgnls types are at the extreme pale end of this co- evident by the habitus illustration ol the original lour spectrum. description (Kamiya 1938a:35). As described by Poppius (1908), there is in ,4. A. kurilensis was described in two different arcticus a geographic variation in the shape of the publications the same year with the lollowing pub- anterior male protarsal claw The three forms re- lication dates (Nakane, in litt.): 1938a - 3l May; cognized by Poppius (1.c.) were later given subspe- 1938b - 5 November. cific rank by Zaitzev (1953:245). The specimens from Mongolia are assigned to the subspecies 1. Acknowledgements. The following persons are thanked for lending me types and other specimens from collec- (Motschulsky), characterized by arcticus alpinus tions under their care: Dr R.B. Angus, Alderhurst, Dr the acute shape of the tooth on the claw. 0. Bistrom, Helsinki, Dr S. Koponen, Tirrku, Dr O.L. Gu6orguiev (1969) described the subgenus Me- Kryshanovskij, Leningrad, Dr G. Scherer, Munich, Dr sogobus for A. insignis. Nilsson et al. (1989) syno- A. Slipinzi, Warszaw, and Dr S. Thgaki, Sapporo. Dr LM. Kerzhner, Leningrad, and Dr T. Nakane, Chiba, based on nymized Mesogabus with Gaurodyles, kindly provided valuable information. Dr D.J. Larson, the synonymy presented above. St. John's, has contributed with valuable comments and ideas.

The Agabus tristis group References Zimmermann (1934) named this group after A. bi- pustulatus (Linnaeus), but Larson (1989) instead Aub6, C. 1838. Hydrocanthares et Cyriniens. In: Dejean, P.E Species g6n6ral des Col6optbres- de la single Holarctic species of based the name on the collection de m. le Comte Dejean. Vol. 6. Paris. the group. Larson (1989) recognized only six spe- Bistrdm, O. 1987. Lists of the types in the Zoolo- cies in this group. It seems likely that also A. adus- gical Museum, University of Helsinki. 3. Coleopte- 160160 Anders N. NilssonNi/sson

generic ra: Haliplidae,Haliplidae, Dytiscidae, Gyrinidae.Gyrinidae. - - ActaActa ent.ent. of the tribetribe Agabini,Agabini, withwith somesome generic sy-sy- fenn.fenn.48:33-40. 48:33-40. nonymy (Coleoptera: Dytiscidae). Dytiscidae). - - Ann. ent.ent. Brinck, P. 1943.1943. Insecta,Insecta, exex SibiriaSibiria meridionali et Mon- Soc. Amer. 35:355-362.35:355-362. golia, inin itinereitinere Orjan OlsenOlsen 1914l9l4 collecta. A. Cole- Motschulsky, V. de. 1853.1853. Hydrocanthares de la Russie.Russie. optera, aa FritzFritz JensenJensen lecta.lecta. VIII.VIII. Haliplidae,Haliplidae, Dy-Dy- Helsingfors. tiscidae, Gyrinidae.Gyrinidae. - NorskNorsk ent. ent. Tidsskr. Tidsskr. Motschulsky, V.V. de. 1859.1859. CokopkresCol6optdres dudu gouverne- gouverne- 6:154-161. ment de Iakoutsk,lakoutsk, recuillies par par M. M. Pavlofski. Pavlofski. - - Gtkorguiev,Gudorguiev, V.V.B. B. 1968. 82.82. Coleoptera: Haliplidae, Dy-Dy- Bull. Phys. Mat. Acad.Acad. St.St. Petersb.Petersb. 17:539-544.17:539-544. tiscidae,tiscidae, GyrinidaeGyrinidae II.II. ErgebnisseErgebnisse desdes zoologischenzoologischen Motschulsky, V. de.de. 1860. Col6optdres Cokopkres de laIa SiUrieSibdrie ori- Forschungen von Dr. Z. KaszabKaszab inin derder Mongolei.Mongolei. entale et en particulier desdes rivesrives de de l'Amour. l'Amour. - In: - Izv. zool. Inst., Sof. 27:23-29. Schrenck, L. von.von. Reisen Reisen undund Forschungen Forschungen- imim Gtkorguiev,Gu6orguiev,- V.B.V.B. 1969.1969. DytiscidaeDytiscidae etet GyrinidaeGyrinidae (Cole-(Cole- Amur-Lande. Vol.Yol. 2:2. St. Petersburg. optera) de Mongolie.Mongolie. - AnnlsAnnls zool. zool. Warsz. Warsz. Nakane, T.T. 1989.1989. The beetlesbeetles of JapanJapan (new(new series)series) 87.87. 27:59-64. - - NatureNature and and Insects 24:18-24.24:18-24. Gtkorguiev,Gu6orguiev, V.V.B. B. 1972.1972.235. 235. Haliplidae, Dytiscidae, Gy- Nilsson,- A.N. 1984.1984. NomenclaturalNomenclatural and distributionaldistributional rinidae IV.IV. Ergebnisse der zoologischen Forschung- notes on some northern speciesspecies of Agabus (Coleop- en von Dr. Z. Kaszab in der Mongolei (Coleoptera). tera: Dytiscidae).Dytiscidae). - - NotulaeNotulae ent.ent. 64:185-188.64:185-188. - Faunist. Abh. staatl.staatl. Mus. Mus. Tierk. Tierk. Dresden Dresden Nilsson, A.N. 1985.1985. NotesNotes onon thethe Agabus congener 4(6):31-44.4(6):3Fa4.- groupqroup (Coleoptera,(Coleoptera, Dytiscidae)Dytiscidae) in KilpisjärviKilpisjiirvi andand Guignot, F.F. 1956a. 1956a. DiagnosesDiagnoses provisoiresprovisoires dede nouveauxnouveaux Aland. - NotulaeNotulae ent.ent. 65:137-141. 65:137-141. dytiscides dudu Magyar NemzetiNemzeti Muzeum. Muzeum. - Bull.Bull. Nilsson, A.N. & Larson, D.J. 1990.1990. AA reviewreview of thethe mens. Soc. Linn.Linn. LyonLyon 25:139.25:139. - Agabus affinisaffinis group group (Coleoptera: (Coleoptera: Dytiscidae), Dytiscidae), Guignot, F.E 1956b.1956b. Dytiscidae nouveaux ou int6ressantsintftessants with the descriptiondescription of aa newnew speciesspecies fromfrom SiberiaSiberia du MuséeMus6e National d'Histoire NaturelleNaturelle dede Buda-Buda- and a proposedproposed phylogeny. - Syst.Syst. Ent.Ent. 15:227-15:227- pest. - Ann.Ann. Hist. Hist. nat. nat. Mus. Mus. Hung. Hung. (N.S.) (N.S.) 239. - 7:395-399.'t:395-399.- Nilsson, A.N., Roughley,Roughley, R.E.R.E. && Brancucci,Brancucci, M.M. 1989.1989. A Guignot, EE. 1957.1957. Quarante-troiskmeQuarante-troisidme notenote sursur lesles Hyd-Hyd- review of the genus- and family-group namesnarres of the rocanthares (Col.).(Col.). -- Bull.Bull. Soc.Soc. ent.ent. Fr.Fr.62:91-94. 62:91-94. family Dytiscidae LeachLeach (Coleoptera). (Coleoptera). - - Ent.Ent. Kamiya, K. 1938a.1938a. A systematic study of the JapaneseJapanese scand.scand. 20:287-316.20:287-316. Dytiscidae. - J.J. Tokyo Tokyo NogyoNogyo Daigaku 5:l-685:1-68 + Paykull, G.G. von.von. 1798. 1798. FaunaFauna Suecica.Suecica. Insecta.Insecta. Vol.Vol. 1.l 7 pls. - Upsaliae. Kamiya, K. 1938b. FamilyFamily Dytiscidae. Dytiscidae. - FaunaFauna Nippo-Nippo- Poppius, B. 1905.1905. Beiträge Beitriige zur zur Kenntnis Kenntnis derder nica 10(8:11):74-87.l0(8: I 1):74-87. - Coleopteren-Fauna des Lena-ThalesLena-Thales inin Ost-Sibi-Ost-Sibi- Kirby, W. 1837.1837. The The insects. insects. - In: Richardson, J. Fau-Fau- rien. 1.I. Haliplidae und Dytiscidae. - Öfv.Ofv. FinskaFinska na Boreali-Americana.Boreali-Americana. PartPart- 4. London. Vet.-Soc. Forhandl.Förhandl. 47(16):1-24.47 (16):1-24. - Köno,K6no, H. 1944.l9zl4. DieDie Käfer-FaunaKiifer-Fauna der Nord-KurilenNord-Kurilen (XI. Poppius, B.B. 1908. 1908. OberUber einigeeinige VariationserscheinungenVariationserscheinungen Beitrag zur Kenntnis der KäferfaunaKiiferfauna derder Kurilen).Kurilen). imim männlichenmiinnlichen GeschlechtGeschlecht bei AgabusAgabus (Gaurody-(Gaurody- - Cishima Gakujutsu-chösa-kentyil-taiCakujutsu-ch6sa-kenty0-tai HökushoH6kusho tes)tes) arcticus Payk.Payk. - Medd.Medd. Soc.Soc. FaunaFauna fl.fl. fenn.fenn. 1:74-92.-1:74-92. 34:52-55.34:52-55. - Lafer, G.Sh.C.Sh. 1988.1988. NovieNovie bidi zjukov-plavuntsov (Cole-(Cole- Poppius, B.B. 1909.1909. Die Coleopteren-Fauna der Halbinsel optera, Dytiscidae)Dytiscidae) s DalnegoDalnego VostokaVostoka SSSR. Pp. Kanin. - ActaActa Soc. Soc. FaunaFauna Fl.Fl. Fenn.Fenn. 31(8):1-583l(8):l-58 + 52-59.52-59. -- In:In: Fauna, Fauna, sistematika sistematika i ibiologija biologija presno-presno- 1I map. vodnich bespozvonotjiych. VladivostokVladivostok (Akademi-(Akademi- Reitter, E. 1899.1899. Beitrag zur Coleopteren-Fauna des rus- ja NaukNauk SSSR).SSSR). sischensischen Reiches und und derder angrenzendenangrenzenden Liinder. Länder. - - Lafer, C.Sh.G.Sh. 1989. Sem. Sem. Dytiscidae Dytiscidae - - Plavuntji.Plavuntji. - Deutsche ent. Zeitschr.Zeitschr. 1899:193-199.1899: 193-199. Opredelitel NasekomichNasekomich DalnegoDalnego VostokaVostoka SSSRSSSR- Sahlberg, J. 1880. 1880. BidragBidrag tilltill nordvestranordvestra SibiriensSibiriens in-in- 3(1):229-253.3(t):229-251. sektfauna.sektfauna. Coleoptera. I.1. - K.K. svenska svenska Vetensk- Vetensk- Larson, D.J.D.J. 1975. 1975. TheThe predaceouspredaceous waterwater beetlesbeetles (Cole-(Cole- Akad. Handl. 17(4):1-115l7(4):l-l15 - + 1I pl. optera: Dytiscidae)Dytiscidae) of Alberta: systematics,systematics, natural Scholz, R. 1917.1917. WissenschaftlicheWissenschaftliche Ergebnisse der Bear- history and distribution.distribution. - Quaest.Quaest. Entomol. Entomol. beitung von 0.O. LeonhardsLeonhards Sammlungen.Sammlungen. 9.9. VierterVierter 11:245-498.ll:245-498. - Beitrag zur KenntnisKenntnis undund Verbreitung Verbreitung paläarkti-paliiarkti- Larson, D.J.D.J. 1989. 1989. RevisionRevision of North AmericanAmerican Agabus scherscher WasserkäferWasserkiifer (Dytiscidae). (Dytiscidae). - Ent. Mitt.Mitt. Leach (Coleoptera: Dytiscidae):Dytiscidae): introduction, keykey 6:250-258. - to speciesspecies groups, and classificationclassification of thethe ambi-ambi- Scholz, R. 1920.1920. 5.5. BeitragBeitrag zurzur KenntnisKennrnis undund Verbreit-Verbreit- guus-, tristis-, andand arcticus-groups. - Can.Can. Ent.Ent. ung paläarktischerpaliiarktischer WasserkäferWasserkiifer (Dytiscidae).(Dytiscidae). - 121:861-919.l2l:861-919. - Ent. Bl.Bl. Biol.Biol. Syst.Syst. KäferKifer 16:13-16.16:13-16. - Larson, D.J.D.J. && Nilsson,Nilsson, A.N.A.N. 1985.1985. TheThe HolarcticHolarctic spe-spe- Sharp, D.D. 1882.1882. OnOn aquaticaquatic carnivorouscarnivorous ColeopteraColeoptera or cies of Agabus (sensu lato) Leach (Coleoptera: Dy-Dy- Dytiscidae. - - Scient.Scient. Trans.Tians. R.R. Dubl.Dubl. Soc.Soc. (2)(2) tiscidae). -- Can.Can. Ent. Ent. 117:119-130. ll7:ll9-130. 2:17-1003.2:17-10O3. LeConte,Leconte, J. 1863.1863. List of thetheColeopteraof Coleoptera of North Ame-Ame- Wewalka, G.C. 1986.1986. Ilybius mandsuricus Guignot gehörtgeh0rt rica. - Smithson.Smithson. Misc.Misc. Coli.Coll. 140:1-49.140:l-49. inin die Gattung Agabus (Dytiscidae, Col.). Col.). - Kole-Kole- Leech, H.B. 1938.1938. DescriptionsDescriptions of threethree newnew speciesspecies of opt. RdschauRdschau 58:109-111.58:109-l I l. - Agabus fromfrom HudsonHudson Bay Bay (Coleoptera: (Coleoptera: Zaitzev, P. 1905.1905. Notizen aber[iber WasserkäferWasserkiifer (Coleoptera Dytiscidae). - - Can.Can. Ent.Ent.70:123-12'1 70:123-127. . aquatica). XI-XX. XI-XX. - Russk.Russk. ent. ent. Obozr. Obozr. Leech, H.B. 1942.1942. KeyKey to the NearcticNearctic genera of waterwater 5:211-215. Notes on East Paloearctic Agabus l6l

Zaitzev, P. 1906. Notizen iiber Wasserkiifer (Coleoptera Schwimmkiifer. Wien. ent. Z. 44i165-187. aquatica). XXI-XXX. Russk. ent. Obozr. Zimmermann, A. 1934.- Monographie der paliiarkti- 6:l'70-175. - schen Dytiscidae. V. Colymbetinae (1. Teil). Ko- Zaitzev, P. 1908. Beitrag zur Kenntnis der Wasserklfer leopt. Rdschau 20:.138-214. - von Chinesisch-Centralasien. Ezheg. Zool. Mus. Zimmermann, A. & Gschwendtner, L. 1935. Monogra- 13:417-426. - phie der paliiarktischen Dytiscidae. VI. Colymbeti- Zaitzev, P. 1910. Beitriige zur Kenntnis der Wasserkiifer nae (2. Teil). Koleopt. Rdschau 2l:61-92. des Ostens von Nordsibirien. Haliplidae, Dytisci- - dae, Gyrinidae, Hydrophilidae, Georyssidae, Dryo- pidae u. Heteroceridae. Zap.lmp. Akad. Nauk. Sammanfattning (8) l8(9):11-52 + pl. lB.- Zaitzev, P. 1913. Neue kaukasische Wasserkiifer. I. - En undersdkning av material av flera diligt ktinda Mitt. Kaukas. Mus. 7:195-199. Ostpalearktiska arter av dykarsliiktet Agabus har ZaiLzev, F.A. 1927. Plavuntsi Kavkaza (Coleoptera, Dy- resulterat i flera nya synonymer. Agobus inexspec- tiscidae). Raboti Severo-Kavkaz. Gidrobiol. Stantsii 2(l):l-42.- tatus sp. n. beskrivs frdn Sibirien. Viktiga karak- Zaitzev, EA. 1953. Nasekomye zhestkokrylye. Plavunt- tArer presenteras for flera arter och lektotyper ut- sovye i vertyachki. - Fauna SSSR 58:l-376. ses. Bestamningsnycklar ges fdr A. labiatus-grup- Zimmermann, A. 1919. Die Schwimmkiifer des Deut- pens alla arter och A. confinis-gruppens palearkt- schen Entomologischen Museums in Berlin- Dahlem. Arch. Naturgesch. 83(1917):68-249. iska arter. Zimmermann,- A. 1928. Neuer Beitrag zur Kenntnis der

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