ANNALES ZOOLOGICI (Warszawa),2005, 55(1): 83-94 A REVIEW OF THE ANTS OF THE GENUS LASIUS FABRICIUS, 1804, SUBGENUS DENDROLASIUS RUZSKY, 1912 (HYMENOPTERA: FORMICIDAE) FROM EAST PALAEARCTIC Alexander Radchenko Museum andInstitute ofZoology, PolishAcademy ofSciences, 64, Wilcza str., 00-679, Warsaw; e-mail:[email protected] — Abstract. The taxonomyofthe antsubgenus Dendrolasius Ruzsky, 1912 is reviewed on the base ofthe investigation oftypes and ofnon-type material ofseveral species. L.fuji is describedas anewspecies, which includes formerL.fuliginosus (LatreiUe, 1798) fromthe EastPalaearctic.L.nipponensisForel,1912,proposedbyEspadaleretal.2001 asthereplace- mentnamefor"oriental^%inoiU5",actuallyisagoodspeciesandtheseniorsynonymofi. crispusWilson,1955;I.orientalisKarawajew,1912isrevivedfromsynonymyandisconside- redastheseniorsynonymofI. teranishiiWheeler, 1928;I. capitatusKusnetzov-Ugamsky, 1928 is considered as a good species, different from L. crispus. A key to Dendrolasius workersandqueensfromtheEasternPalaearcticisalsogiven. — Key words. Ants, taxonomy, Lasius, Dendrolasius, new species, new synonyms. East Palaearctic. Introduction Forel, 1912 from Japan, and L.fuliginosusvar. orientalis Karawajew, 1912 from Korea. A little earlier Wheeler The ants ofthe subgenus Dendrolasius Ruzsky, 1912 (1910),basedonthesingleverypeculiarqueenfoundin are the most peculiar both morphologically and bio- Japan, describedI. spathepus, which alsobelongs to the logicallyamongallLasiusFabricius, 1804 species.These subgenusDendrolasius. so-called jet black ants have a quite high level ofsocial Later on, several Dendrolasius species and infra- organisation, formlarge colonies, usuallybuild big car- specific forms were described: Acanthomyops fuligi- ton nests in living trees, and are characterised by their nosus var. capitatus Kusnetzov-Ugamsky, 1928 from specific, strong smell. All ofthem are distinctly bigger RussianFarEast;L. teranishiiWheeler, 1928fromJapan; than other Lasius species (workers' body size up to 4.5 L. ouchiiTeranishi, 1940fromJapan;L. buccatusStarcke, - 5.5 mm), have shiny black or dark brownish-black 1942 from Bosnia; I. crispus Wilson, 1955 from Japan body,andadistinctlyemarginateoccipitalmarginofthe andKorea; L. morisitaiYamauchi, 1978 from Japan. headinmostspecies. After complete or local revisions and reviews of This subgenus includes eight species only (taking the genus Lasius or its subgenus Dendrolasius (Wilson into accountthepresentrevision), two ofwhichdistrib- 1955; Yamauchi and Hayashida 1968; Yamauchi 1978; uted in the Western Palaearctic (as far east as the Altai Kupyanskaya 1989, 1990; Espadaler et al. 2001; see also Mts.), and six others found in the eastern part of the Bolton 1995) the number ofthevalidDendrolasius spe- Region, e. g. in thesouthernpartofRussian FarEast (as cies was reducedto seven. far west as the Amursky Region), north-eastern China, Despite the presence of many pubhcations dealing KoreanPeninsula,andJapan;onespecieswasalsofound with the subgenus Dendrolasius, there were still quite a inTaiwan. lotofunresolvedtaxonomic questions. Ihadthe oppor- Ruzsky (1912) described the subgenus Dendrolasius tunity to investigate the types of several species and including one species - L. fuliginosus (LatreilTe, 1798). infraspecific forms, what allowed to me to solve some In the same year, two new forms belonging to this sub- problems and clarify the taxonomic situation of this genus were described: L. fuliginosus var. nipponensis Palaearctic subgenus. . 84 A. Radchenko Material and methods "v. nipponensis Forel, worker, Type, Tokyo" (both hand written by Forel), "Coll. Forel", "Typus" (red printed This work is based on the investigation of some label) (MHNG); "paralectotype" queen, "L. fuliginosus collected personally material and oftype and non-type LatreiUe, $, Japan (Yano)", "v. nipponensis Forel, $, specimens, preservedin different museums andinstitu- Type, Tokyo" (bothhandwrittenbyForel), "Coll. Forel", tions: Museiun ofthe Comparative ZoologyofHarvard "Typus" (red printed label) (MHNG) (really this queen University, Cambridge, Massachusetts, USA (MCZ); is L.fuji, see below). Non-type material: more than 20 Museum d'Histoire naturelle, Geneva, Switzerland workers, 2 queens from the Russian Far East, North (MHNG);InstituteofZoologyoftheUkrainianNational Korea andJapan (IZK, MIZ, ZMMU, BPI, JUK). AcademyofSciences,Kiev,Ukraine(IZK);Museimiand Measurements and indices of the paralectotype Institute ofZoology ofthe Polish AcademyofSciences, workers: HLi = 1.09-1.25, HL2 ^ 1.15-1.30, HWi = Warsaw, Poland (MIZ); Jagiellonian University, Cracow, 1.09-1.26,HW2=0.67-0.76,SL= 1.01-1.18,OL=0.23- Poland (JUK); ZoologicalMuseum ofthe MoscowState 0.25, AL = 1.26-1.51; CI = 1.00-1.01, CLI = 1.04-1.05, University, Russia (ZMMU); Zoological Institute ofthe CWI = 1.63-1.67, SIi = 0.92-0.94, SI2= 0.92-0.94, 01 = Russian Academy of Sciences, St.-Petersburg, Russia 0.20-0.21. (ZIN); Biological and Soil Institute of the Russian Diagnosis ofworkers andqueens. Workers: petiolar AcademyofSciences, Vladivostok, Russia (BPI). scale (seen in profile) thin, distinctly narrowing to the The following measurements (in mm) and indices top and with flattened dorsal crest, symmetrical; seen are used: in front or from behind, it is the vwdest at the level of HLi - head length in fiiU-face view, measured from the spiracles, clearly tapering to the top, its dorsal crest the mid-point ofthe occipital margin to the mid-point narrowly rounded, without notch; head in full face oftheanteriorclypealmargin;HL2-headlengthinfull- view distinctlynarrowing anteriorlyand with shallowly face view, measured as a diagonal Une from the upper- emarginate occipital margin; scape, midand hindtibiae mostpointofanoccipitalcornertothemid-pointofthe not flattened, elliptical in cross-section; ratio of min/ anterior clypeal margin; HWi - maximum head width max diameters of the scape > 0.7; scape and legs with, infull-faceview, excludingeyes; HW2 - minimumhead numerous subdecimibent hairs; promesonotal dorsum width near the level of tiie mandibular insertion; SL and occipital niargin with rather long standing hairs; - maximum straight-line length ofthe antennal scape bodyvwth sparse, shortdecumbentpubescence. in profile; OL - maximum diameter of the eye; AL - Queens: petiolar scale (seen in profile) thin, narrow- diagonal length ofthe alitrunkseen in profile, from the ingto the top;headin fuUfaceview distinctlynarrowing neck shield to the posterior margin ofpropodeal lobes anteriorly,withalmoststraightlateralmarginsinfrontof (workers), or from the anterio-dorsal point ofalitrunk the eyes, and with hardly emarginate occipital margin; toposteriormargin ofpropodeallobes (queens). scape, mid and hind tibiae not flattened, elliptical in Indices: cross-section; ratio ofmin/max diameters ofthe scape > CI = HWi / HLi; CLI = HL2 / HLi; CWI = HWi / 0.7;legsandscapewithdensedecumbentpubescenceand HW2; SIi = SL/HLi; SI2 = SL/HWi; 01 = OL / HWi. numeroussubdecumbenthairs;head, alitrunkand gaster withveryabimdant,long, often curvedstandinghairs. Notes. Forel (1912) described L. fuliginosus var. nip- Taxonomy ponensis from workers and queen from Japan (vicinity of Tokyo). The most important features, separating the Lasius (Dendrolasius) nipponensisForel, 1912, stat. rev. workers of this form from those of L. fuliginosus, have (Figs 1-7, 15-21) beenemphasisedintheoriginaldescription: "... lecaillede Vouvriere estassez tranchante et non obtuse, la tete estplus Lasiusfuliginosusvar. nipponensisForel 1912:339,w,neeq,Japan. retrecie devant. ." [e.g.: ...thepetiolarscaleoftheworkeris Lasiusnipponensis:Santschi 1941:278 (raisedtospecies). rathersharp(thin)andnotblunt, theheadisnarrowerante- L.fuliginosus var. nipponensis: as junior synonym ofL. fuliginosus riorly...] (loc. cit., p. 339). Thesefeatures fullycorrespond (LatreiUe):Wilson 1955: 138;Yamauchi 1978: 171; Bolton 1995: tothoseofthetypespecimens (see above). 224;revivedfromsynonymyandraisedtospeciesasthefirstava- The taxonomic position of L. nipponensis is some- ilablereplacementnamefor"ohentalfuliginosus":Espadaleretal. what confused because the queen, described by Forel Lasiu2s00c1r:is3p4u0s;WIimlasione,ta1l9.5250:031:446,0.w,q,m,Japan(unresolvedprimary (1912), indeed belongs to the "oriental /u%«05W5". It homonym of Lasius crispus Theobald, 1935: 68, France, Mio- has a thick, shghdy narrowing to the top petiolar scale Pliocene); Yamauchi and Hayashida 1968: 401, 402; Yamauchi (seen in profile); mid and hind tibiae and tarsi with 1978: 174,syn.nov. dense decumbent pubescence only, without subdecum- bent hairs; the head in full face view only slightly nar- Materialexamined.Paralectotypes, 3workersonthe rowing anteriorly, with distinctly emarginate occipital same pin, "L. fuliginosus Latr., worker, Tokyo (Yano)", margin, and with abundant, though relatively short. SUBGENUS DENDROLASIUS RUZSKY (FORMICIDAE) FROM EAST PALAEARCTIC 85 and molecular evidence that ants from the eastern Palaearctic, previ- ously determined as L. fuliginosus, are not conspecific with the "typi- cal"L.fuliginosus (Latreille), distrib- uted in the western Palaearctic (e.g. as far east as the Altai Mts.), and i proposed for this species the name L. nipponensis Forel (as the oldest nameattributedto"oriental/uZi^mo- sus"). Yet, they did not examine the typesofvar. nipponensis. Actually,L. nipponensis is not conspecific with u "oriental^/igiMosus",whoseworkers have a much thicker, not narrowing atthetoppetiolarscale (seeninpro- file),alessanteriorlynarrowinghead with distincdy emarginate occipi- tal margin, etc. (compare Figs 1-7 and 52-58). Furthermore, queens of these species differ even more than theirworkers (see above). Wilson (1955) described L. cris- pusfromworkers, queensandmales from Japan and Korea. The most iinportant diagnostic features of the workers were: "...femora with numerous outstandingdecumbent to suberect hairs. Cephalic and gastric pilosity denser than in fuliginosus. Petiolar crest viewed from the side thinnerandsharper than in L.fulig- inosus, the anterior and posterior 14 faces less convex..." {loc. cit, p. 144). According to this and the detailed Figures 1-14. 1-7. Lasius nipponensis Forel, 1912 (worker, paralectotype) and 8-14 L.fuji sp. nov. treatment of L. crispus given by ("paralectotype" queen ofL. nipponensis); (1, 8) head, frontal view; (2, 9) alitrunk and petiole, in Yamauchi (1978) and Kupyanskaya profile;(3, 10)petiolarscale,frontalview;(4, 11)scape,dorsalview;(5, 12)same,lateralview;(6,13) (1989, 1990, as junior synonym of hindtibiaandfirsttarsaljoint,lateralview;(7, 14)same,dorsalview.Scalebar= 1 mm. L. capitatus), it is obvious that L crispus has the same diagnostic fea- Standing hairs projecting from its outline (see Figs tures ofL. nipponensis. Firstofall, the thinpetiolarscale, 8-14). All these features correspond with the current the distinctly anteriorly narrowing head, tiie hairy legs treatmentofthequeens ofthe"oriental/w%mo5us" (see and scape, etc., combined with the not flattened scape; also Yamauchi and Hayashida 1968; Yamauchi 1978; alltheseledmeto formallysynonymiseL. crispuswithL. Kupyanskaya 1989, 1990, andFigs 59-65). nipponensis. Wilson (1955) synonymised this name with Lfuligi- Distribution. Southern part of Russian Far East nosus, and wrote such comments about the lectotype (PrimorskyRegion),KoreanPeninsula,Japan(Hokkaido, specimen ofvar. nipponensis, designated by him: "... a Honshu, Shikoku), Taiwan. worker in the Forel collection. Head and thorax partly crushed and not measurable. Pilosity ofpetiolar dorsal margin long, characteristic of the Japanese population Lasius capitatus (Kusnetzov-Ugamsky 1928) alreadydescribed..'' {loc. cit.,p. 143). Most ofsubsequent (Figs 22-28) authors (YamauchiandHayashida 1968; Yamauchi 1978; Kupyanskaya 1989, 1990; Bolton 1995) reputed this Acanthomyopsfuliginosussubsp. capitatusKusnetzov-Ugamsky1928: 18, synonymy without any further explanations. However, w,RussianFarEast(Kusnetzov-Ugamsky^1927: 187,nomennudum); Espadaler et al. (2001) provided both morphological transferredtoLasiusasthejuniorsynonymofLJiiliginosus(LatreiEe): 86 A. Radchenko Wilson 1955; 138;revivedfromsynonymyandraisedto speciesas theirpetiolar scale is somewhatthickerthan in Fig. 4 of theseniorsynonymofI.crispusWilson:Kupyanskaya1989:785,w, Kusnetzov-Ugamsky (1928). q,m;Kupyanskaya1990:229;Bolton1995:222("Turkestan"iserro- In my opinion, for several reasons L. capitatus had neouslygivenasthetypelocality);Imaietal.2003:60. the most ambiguous taxonomic position among all Material examined. Lectotype worker, "Okeanskaya Dendrolasius species. First of all, even the usage of the nr.Vladivostok, Siberia, VIII-1926,N. Kusnetzov" (prob- name "capitatus" was misleading. It was introduced by ablyhand written byWilson); "Lasiusfuliginosus capita- Kusnetzov-Ugamsky (1927) as nomen nudum. He wrote tus Kusnetzov LECTOTYPE" (hand written byWilson); "...Generally, thisspecies [e. g.L.fuliginosus] is verystable "M.C.Z.Type30143"(MCZ,designatedbyWilson,1955); on its characters; only the separate subspecies, recently paralectotype worker, "Okeanskaya nr. Vladivostok, described by me, A. fuliginosus capitatus, nov. (manu- Siberia, VIII-1926, N. Kusnetzov" (hand written by script), lives in theFarEast(South UssuriRegion)..." {loc. Wilson); "Turkestan N. Kusnetzov" (Sic!) (printed origi- cit.,p. 187). Laterhehasprimarilyusedforthis form the nal label from Kusnetzovs collection); "Lectotype nest name "Acanthomyops fuliginosus orientalis Karawajew" series" (hand written by Wilson); "Cotype" (red printed and noted "...all specimens of Acanth. (Dendrolasius) label); "Lasius fuliginosus capitatus Kuzn.-Ugams. det. fuliginosus Latr., collected by me in Ussuri Region [now E. O. Wilson" (hand written by Wilson); "M.C.Z. Type - Primorsky Region of Russia], belong to the separate 30143"; paralectotypeworker, "Acanthomyopsfuliginosus taxonomic unit, which Karawajew described as separate orientalis (= capitatus m.) Far East Station Okeanskaya subspecies. Discriminatingfeatures ofthis subspecies are: nearVladivostok" (probablyhandwrittenbyKusnetzov); maxillarypalps rather long 6-jointed, their thirdjoint is "Turkestan N. Kusnetzov" (Sic!) (original printed label the longest, each subsequentjoint is shorter than thepre- from Kusnetzov's collection); "Lectotype nest series" ceding one...Head without emargination on the occipital (hand written by Wilson); "Cotype" (red printed label); margin, or, at most, very shallowly emarginate,... with "M.C.Z. Type30143" (MCZ).Non-typematerial:5work- broadly rounded occipital corners...Petiolar scale "flatty- ersfromthe RussianFarEast (IZK, BPI). rounded" [I do not understand properlywhat the author Labels of paralectotype specimens are misleading correctly meant even in Russian] ...The main features for the presence ofthelabels "Turkestan N. Kusnetzov" [e.g. separatingthis form from Lfuliginosus] are: lack of together with the locality "Far East Station Okeanskaya the emargination on the occipital margin ofthe head and nearVladivostok". Probablyitis simplyalabellingerror a differentstructure ofthe maxillarypalps..." (Kusnetzov- in Kusnetzovs collection. Ugamsky 1928, p. 17). Then, on p. 18 ofthat paper, the Measurements and indices of the type specimens author imexpectedly used for this subspecies the name (data ofthe lectotype are in brackets): HLi = 1.25-1.29 "Ac. capitatus"; he also used the same name (as "subsp. f. (1.29), HL2= 1.32-1.36 (1.36), HWi = 1.22-1.27 (1.27), nov.") in the legend for Figs 1-4. In the following paper HWa = 0.78-0.85 (0.85), SL = 1.11-1.12 (1.12), OL = (Kusnetzov-Ugamsky, 1929) he used for the same Far 0.24-0.27 (0.27), AL = 1.34-1.44 (1.44); CI = 0.97-1.00 Eastern form of I. fuliginosus the Karawajew's name (0.99), CLI = 1.05-1.07 (1.05), CWI = 1.49-1.59 (1.49), "orientalis". Formally, the name "capitatus" couldbe con- SIi = 0.87-0.89 (0.87), SI2 = 0.89-0.92 (0.88), 01 = sidered as an unnecessarilyproposed replacement name L 0.19-0.21 (0.21). forvar. orientalis, but in fact, orientalisis another spe- Queens andmales are unknown. cies,thatdiffersbothfrom"oriental^%f«05Ms"andfrom Diagnosis. Workers: petiolar scale (seen in profile) capitatus (seebelow). relativelythin, though not flattened at the top, approxi- For manyyears thetaxonomic status ofthe subspe- mately inversely V-shaped; when seen in front or from cies capitatus was enigmatic, until Wilson (1955) des- behind, itis slightlynarrowing, not tapering to the dor- ignatedthelectotype ofthisformandconsidereditasa sal crest; head with convex sides, gradually and slightly junior synonym ofI.fuliginosus with such comments: narrowing anteriorly, with rather shallowly emargin- "[the lectotype]...Possessm^ a shallow occipital emar- ate occipital margin; scape, mid and hind tibiae not gination and shortpetiolar hairs, both ofwhich charac- flattened, elliptical in cross-section; ratio of min/max ters seem topredominate in north-eastern Asm..." (loc. diameters ofthescape > 0.7; scapeandlegswithdecum- cit., p. 143). bent pilosity only; promesonotal dorsum and occipital On the contrary, Kupyanskaya (1989), according to margin with short and sparse standing hairs; thirdjoint Kusnetzov-Ugamskysoriginaldescriptionanddrawings ofmaxillarypalps is the longest, each followingjoint is (see above) believed that the main diagnostic features somewhatlongerthanthe nextone; Thecombinationof of L. capitatus and L. crispus are actually the same. these features distinctly separates this species both from Hence, she revived the name capitatus from synonymy, I. nipponensis (see above and Figs 1-7) and from L.fuji raised it to species rankas senior synonym ofL. crispus (seebelowandFigs 52-58). Wilson and provided some details characteristic of all Notes. The type specimens of var. capitatus gener- of the three castes, although she considered the types ally correspond to the original description, except that ofL. capitatus as probablylost. However, Kupyanskaya's SUBGENUS DENDROLASm RUZSKY (FORMICIDAE) FROM EAST PALAEARCTIC 87 Lasius orientalisKarawajew, 1912, Stat, rev., stat. nov. (Figs 29-37) Lasiusfuliginosusvar. orientalis Karawajew, 1912: 586, w, Korea; Wheeler 1928: 123;asjuniorsynonymofL.fuliginosus (Latreille):Wilson 1955: 138;Yamauchi 1978:171;Kupyanskaya1989:783;1990: ^28; Bolton 1995: 225; asjunior syno- nym ofL. nipponensis Forel: Espadaler et.al.2001:341;Imaietal.2003:60. Lasius (Chthonolasius) teranishii Wheeler, 1928: 120, q, Japan, nom. nov. pro Lasius umbratusr. Teranishi 1927: 92, nee Nyiander 1846: 1048 etauct.;transferred to subgenus Dendrolasius: Wilson 1955: 146;YamauchiandHayashida 1968:399, w,m;Yamauchi 1978: 174; Kupyanskaya 1989: 786; 1990: 230; Bolton 1995: 226; ImaietaL2003:59,syn.nov. Lasius (Chthonolasius) ouchii Teranishi, 1940:76,q,Japan;asjuniorsynonymof L.teranishii:Wilson1955:146,syn.nov. (Ihaveneverseenanymaterialreferred to I. ouchii and this new synonymy is established following Wilson's, 1955 objectivesynonymyforI. teranishii). Material examined. Neotype worker (designated here, see below), "Buhta Gaidamak, No. 3196, 21.V.1900, leg. R Shmidt", "Lasius fuliginosus var. orienta- lis Karaw. Typus" (both labels hand writtenbyKarawajew) (IZK). Non- type material: 16 workers from the nest ofneotype; more than 20 workers firom the Russian Far East, Figures 15-28. 15-21 Lasius nipponensis Forel, 1912 (queen) and 22-28 I. capitatus Kusnetzov- North Korea andJapan (IZK, MIZ, Ugamsky,1928(worker,lectotype).(15,22)Head,frontalview;(16,23)alitrunkandpetiole,inprofile; ZMMU, BPI). (17,24)petiolarscale,frontalview;(18,25)scape,dorsalview;(19,26)same,lateralview;(20,27)hind Measurements and indices of tibiaandfirsttarsaljoint,lateralview;(21,28)same,dorsalview.Scalebar= 1mm. the neotype of I. orientalis (data are inbrackets) and oftheworkers (1989, 1990) "I. capitatus"infactreferstoL. nipponensis from the nest ofthe neotype specimen (n - 16) [mean (and itsjunior synonymL. crispus, see above) and does data are in square brackets]: HLi = 1.06-1.26 (1.22) notmatchwiththe type specimens ofL. capitatus. [1.17], HL2= 1.12-1.33 (1.32) [1.25], HWi = 1.04-1.23 L. capitatus seems the most similar to L. morisitai by (1.19) [1.15], HW2 = 0.70-0.80 (0.77) [0.75], SL = the shape ofthe petiolar scale and bythe short standing 1.01-1.12 (1.09) [1.06], OL=0.18-0.27 (0.25) [0.23],AL hairs on alitrunkandhead, butworkers ofthe latterspe- = 1.18-1.44 (1.34) [1.32]; CI = 0.96-1.00 (0.98) [0.98], cies have a somewhat transversal head, which is wider CLI = 1.05-1.09 (1.08) [1.07], CWI = 1.49-1.57 (1.55) than long (CI < 1.00 in type specimens ofL. capitatus [1.54], SIi = 0.88-0.95 (0.90) [0.90], SI2 = 0.89-0.96 and > 1.04 in L. morisitai); these species also differ by (0.92) [0.92], 01 = 0.18-0.22 (0.21) [0.20]. thepresence ofshortbutdistinctsubdecumbenthairson Diagnosisofworkers andqueens. Workers: petiolar the scape ofL. capitatus, while scape ofL. morisitai has scale (seen in profile) thick, low, not narrowing to the dense decumbent pubescence only (compare Figs 25, 26 top; when seen in front or from behind, it gradually and69, 70). f widens to the top; scape distinctly flattened, ratio of Distribution. Southern part of Russian Far East min/max diameters ofthe scape < 0.5; head with con- (PrimorskyRegion). vex sides, gradually and slightly narrowing anteriorly. A. Radchenko species. Furthermore, 17 work- f\ ers among this material, collected lliH in Primorsky Region of Russia ("Buhta Gaidamak, No. 3196, 21.V.1900, leg. RShmidt"), bear also the additional Karawajew's label "Lasius fuliginosus var. ori- entalis Karaw. Typus". However, theycannotformallybelongtothe r type series, because in the original description another type locality and date ofcollecting were given. 'i 5 I do notknowwhy Karawajewdid 34 35 ^ *"^^ ^ mistake, especially because \i he usually worked very accurately and left his own collection for the next generations of myrmecolo- gistsinperfectconditions.Though types ofvar. orientalis seem lost, I believe that specimens, collected about in the same region as the types, and originally determined by Karawajew as the species he described, really belong to var. orientalis. Hence, I designate as the neotype of L.fuliginosus var. orientalis Karawajew the worker Figures29-37.29-35LasiusorientalisKarawajew,1912.(worker,neotype;36,37,queen[afterYamauchi from "Buhta Gaidamak, No. 3196, and Hayashida 1968]); (29, 36) head, frontalview; (30) alitrunkandpetiole, in profile; (31) petiolar 21.V.1900, leg. R Shmidt", labelled scale,frontalview;(32)scape,dorsalview;(33)same,lateralview;(34)hindtibiaandfirsttarsaljoint, byKarawajew as "Typus". lateralview;(35)same,dorsalview;(37)bodyinprofile.Scalebar= 1 mm. Wilson(1955) synonymisedvar. orientalis with L.fuliginosus with with emarginate occipital margin; scape and legs with such comments: . Since the types are not available, numerous short subdecumbent hairs; promesonotal synonymy in this case is tentative. The differences stated dorsiun and occipital margin with a few quite short in the original description are ofa trivial nature, and it standinghairs; would seem that ifKarawajew had really had spathepus Queens: petiolarscale (seeninprofile) thick,low, not beforehim insteadoffuliginosus, hewouldhavenoticedat narrowingtothetop;when seenin frontorfrombehind leastoneoftheseveralexcellentcharacterswhichseparate it graduallywidens to the top; scape and legs, including workers ofthese two species.." (loc. cit., p. 143), though the first tarsaljoint, remarkablyflattened, ratio ofmin/ he never saw the types of this form. Yamauchi (1978) maxdiameters ofscapeandhindtibiae < 0.4; headwith and Kupyanskaya (1989, 1990) repeated this Wilson's emarginate occipital margin, but not cordiform and at synonymy, while Espadaler et al. (2001) consideredvar. most sUghtlywider than long; legs and scape with very orientalis as the junior synonym ofI. nipponensis (see dense decumbent pubescence; head, aUtnmk and gaster Notestothelatterspecies, above). Thename"orientalis" with very dense decumbent pubescence; alitrunk dorsum wasalso usedbyKusnetzov-Ugamsky(1928, 1929);how- withsparse,shorterecthairs. ever, the specimens from Russian Far East, collected and Notes. Karawajew's (1912) description ofL.fuligi- investigatedbyhim,belongto another species (seeNotes nosus var. orientalis (workers) was very short and toL. capitatus, above). insufficient, and the most important diagnostic fea- The most astonishing fact is that var. orienta- tures ofthis formwere notpointed at. The typelocali- lis is neither I. capitatus, nor "orientalfuliginosus", nor ties ofthis species are "Koredschi und Dore, Halbinsel I. nipponensis. All specimens from Karawajew's collec- von Korea (19.VII.1900, P. Schmidt)" (Karawajew tion mentioned above, including the neotype, have the 1912, p. 582). In Karawajew's collection (IZK) I have distincdyflattenedantennalscapeandtheverythick,low found a couple ofworkers from Russian Far East and petiolar scale (seen in profile), which is gradually wid- Korea, determined by Karawajew as L.fuliginosus var. enedto the apexwhen seenin front orfi-ombehind (see orientalis. All ofthem undoubtedlybelongto the same Figs 29-35). These diagnostic features completelymatch SUBGENUS DBNDROLASIUS RUZSKY(FORMICIDAE) FROM EAST PALAEARCTIC 89 thoseofL. teranishiiWheeler(see Yamauchi and Hayashida 1968; Yamauchi 1978; Kupyanskaya 1989, 1990),whatletmeconsider L. orientalis Karawajew, 1912 as the senior synonym ofL. teran- ishiiWheeler, 1928. Distribution. Southern part W of Russian Far East (Amursky and Primorsky Regions, Islands Shikotan and Kunashir), Korean Peninsula, Japan (Hokkaido, northern andcentralHonshu). U LasiusspathepusWheelei, 1910 43 44 (Figs 38-51) Lasius spathepus Wheeler 1910: 130, q, Japan; 1928: 121,w,m;Wilson 1955: 147; Yamauchi and Hayashida 1968: 401;Yamauchi1978:172;Kupyanskaya 1989: 787; 1990: 231; Bolton 1995: 225;ImaietaI.2003:59. Materialexamined.Holotype queen: "Japan, Kuwana coll., 1910", "Type", "Holotype Lasius spathepus Wheeler", "M.C.Z. type 71691" (MCZ). Non-type inaterial: about 30 workers, 1 queen fromthe RussianFarEast, North Korea and Japan (MCZ, IZK, MIZ, ZMMU, BPI, JUK). Measurements andindices of the holotype queen: HLi = 1.66, HL2= 1.85, HWi = 2.03, HW2= 1.28, SL = 1.55, OL = 0.43, AL = mm; 2.53 CI = 1.22, CLI = 1.11, CWI = 1.59, SIi = 0.93, SI2 = 0.76, 01 = 0.21. Diagnosis of workers and queens. Workers: petiolar scale (seen in profile) thin, distinctly narrowing to the top, Figures 38-51. Lasius spathepus Wheeler, 1910 (38-44, worker; 45-51, asymmetrical; when seen in front or from behind it queen,holotype);(38,45)head,frontalview;(39,46)alitrunkandpetiole, inprofile;(40,47)petiolarscale,frontalview;(41,48)scape,dorsalview; graduallynarrowing to the top, with straight or shghtly (42,49)same,lateralview;(43,50)hindtibiaandfirsttarsaljoint,lateral notched dorsal crest; scape distinctly flattened, ratio of view;(44,51)same,dorsalview.Scalebar= 1 mm. min/max diameters ofthe scape < 0.5; head with con- vex sides, gradually and slightly narrowing anteriorly, with emarginate occipital margin; scape and legs with thanlong; scapeandlegs, includingthefirsttarsaljoint, numerous short subdecumbent hairs; promesonotal remarkably flattened, ratio of min/max diameters of dorsum and occipitalmarginoftheheadwithrelatively scape and hind tibiae < 0.4; legs with dense decum- sparse andlong standinghairs; bentto subdecumbentpubescence; antennal scapewith Queens:petiolarscale (seeninprofile) thin,^distinct- abundant subdecumbent to suberect hairs; head, ah- lynarrowingto thetop, asymmetrical; headwithdeeply trunkand gasterwithveryshort and sparse decumbent emarginateoccipitalmargin, cordiform,distinctlywider pubescence, alitrunkdorsumwithoutstandinghairs. . 90 A. Radchenko Notes. Lasius spathepus is the mostpeculiarspecies ofthegenus, especially for the characters ofits queens(seeaboveandFigs45-51). Workers are similar to those ofI. orientalis by their distinctly flat- tened scape, but well differ from the latter by the much thinner, narrowing at the toppetiolar scale (seen in profile, compare Figs 39, 40 and30, 31). On theotherhand, the shape of petiolar scale recalls L. nipponensis, butin thelatterthe scapeisnotflattened. Wheeler (1910) described L. spathepusfromthesinglequeen fromJapan (he describedworkers and males in 1928). This queen has a so peculiar shape of legs and head that Wheeler discussed the possibility ofthe existence of twodifferentmorphsofqueensin thatLasius species. Further inves- tigations showedthatL. spathepus has unusual qUeens only, rather than normal dhes (Wilson 1955; Yamauchi 1978; Kupyanskaya 1989, 1990; Imai et al. 2003). Distribution. Southern part of RussianFarEast(knownonlyfrom one locality: Primorsky Region, Anisimovka), Korean Peninsula, Japan (allfourmainIslands). Lasius morisitaiYamauchi, 1978 (Figs 66-79) Lasius morisitai Yamauchi 1978: 176, w, q, m, Japan; Kupyanskaya 1989: 787; Figures 52-65. Lasiusfuji sp. nov. (52-58,worker, holotype; 59-65, queen, paratype); (52, 59) head, 1990: 232; Bolton 1995: 224; Imai et frontalview;(53,60)alitrunkandpetiole,inprofile;(54,61)petiolarscale,frontalview;(55,62)scape, al.2003:60. dorsalview; (56, 63) same,lateraslavmiee,w;do(r5s7a,l6v4i)ewh.iSncdalteibbiaara=nd1fmirmst.tarsaljoint,lateralview; (58,65) Material examined. About 20 workers, 2 queens pilosity only, without standing hairs; promesonotal fromtheRussian FarEast,NorthKoreaandJapan (IZK, dorsumandoccipitalmarginwithabundant,quiteshort MIZ, ZMMU, BPI, JUK). standinghairs; Diagnosis of workers and queens. Workers: peti- Queens: petiolar scale (seen in profile) is relatively olar scale (seen in profile) relatively thin, though not thin, though not flattens at the top; head with convex flattened at the top, approximately inversely V-shaped; sides, gradually and slightly narrowing anteriorly, and when seen in front or from behind, it is only slightly with shallowly emarginate occipital margin; scape, mid narrowing to the dorsal crest; head shorter than broad and hind tibiae not flattened, elliptical in cross-section; (CI > 1.04), with convex sides, gradually and slightly ratio ofmin/max diameters ofthe scape > 0.7; legs and narrowing anteriorly, and with distinctly emarginate scape with dense decumbent pubescence only; head, occipital margin; scape, mid and hind tibiae not flat- alitrunk and gaster with very sparse short standing tened,ellipticalincross-section;ratioofmin/maxdiam- hairs, and with extremely short and sparse decumbent eters ofthe scape > 0.7; scape and legs with decumbent pubescence, appears shiny. SUBGENUS DENDROLASm RUZSKY (FORMICIDAE) FROM EAST PALAEARCTIC 91 Notes. Lasius morisitai was the last species, described in the subge- nus Dendrolasius (Yamauchi 1978). Its workers mostresemble L. capita- tus, differing from them mainly by the wider, transverse head and by the absence of subdecumbent hairs on the antennal scape; they are also similarto L.fuji and differ from the latter by the wider head, shorter standing hairs on the alitrunk dor- sum and somewhat thinner peti- olar scale (compare Figs 52-58 and 22-28, 66-72). However, queens of I. morisitaiweU differfrom those of I.fujibytheverysparse decumbent pubescence of the body and much shorter and less abundant standing hairs on the alitrunk dorsum (com- pare Figs 59-65 and73-79). Distribution. Southern part of RussianFarEast(PrimorskyRegion, UssuriyskyNaturalReserve),Korean Peninsula, Japan (central Honshu). Asmentionedabove, Espadaleret al. (2001)providedbothmorphologi- cal and molecular evidence that the "oriental^%'«osu5" is adistinctspe- cies,differsfromthe"typical"Lfulig- inosus (Latreille) distributed in the westernpartofthePalaearcticRegion (e.g.asfareastastheAltaiMts.), and have proposed for it the replacement name L. nipponensis Forel (as the oldest name referred to this species). Nevertheless, neither I. nipponen- sis, nor any other name referable to Dendrolasius,cannotbeusedinstead of "oriental fuliginosus", since all of themare assignedtootherspecies. In Figures66-79.LasiusmorisitaiYamauchi,1978(66-72,worker;73-79,queen);(66,73)head,frontal suchasituationIhavetodescribedas view; (67, 74) alitrunkand petiole, inprofile; (68, 75) petiolar scale, frontal view; (69, 76) scape, newthefollowingspecies. dorsalview;(70,77)same,latersalamvei,ewd;or(s71a,l7v8i)ewh.iSncdalteibbiaara=nd1fmirmst.tarsaljoint,lateralview;(72,79) Lasiusfuji sp. nov. Materialexamined. Holotype, worker. North Korea, (Figs 52-65) Prov. Chagang, Myohyang-san Mts., way to Pirobong, No. 275-85, 25.VI.1985, leg. M. Woyciechowski (IZK); Lasiusfuliginosus:easternpopulationsonly,distributedfromAmursky paratypes: 15 workers, 6 queens from the same nest as R13e8g;ioYnamoafuRcuhsisiaantdoHJaapyaans;hirdeavis1i9o6n8s:a3n9d8;reYvaimeawsu:chWiil1s9o7n8:1915751:; tHhweanhgolhoatey-ppeu;kmtoor,e8tkhmanW50Hwaoerjkue,rsS:ujNaonrgt-hsaKnoMrtesa,, P5r5ovm. Kupyanskaya1989:783;1990:227;WuandWang1995:156;Bolton a.s.l, Pinus-Quercus forest. No. 44-85, 15.VI.1985; ibid., 1995:223,neeLatreiUe1798:36. 600ma.s.l,youngQuercusforest.No. 81-85, 17.VI.1985; Lasius nipponensis: asjunior synonym ofL.fuliginosus: Wilson 19W55u: Prov. Chagang, Myohyang-sanMts,belowChonjuRock, 138; Yamauchi 1978: 171; Kupyanskaya 1989: 783; 1990: 227; m etWang 1995: 156;Bolton 1995:224;revivedfromsyijonymyand No. 169-85, 22.VI.1985, 230 a.s.l, deciduous forest; raisedtospeciesasthefirstavailablenamefor"oriental^%'«osMi": Prov. Chagang, Myohyang-san Mts, near monastery Espadaleretal.2001:340;Imaietal.2003:60,neeForel1912:339. Pliodae, No. 217-85, 24.VI.1985, 540 m, a.s.l pine- 92 A. Radchenxo chestnut forest; Prov. Samjion, shore of Lake Samjion, 1.68-1.70 [1.69], SIi = 0.91-0.93 [0.92], SI2= 0.88-0.90 No. 596-85, 10.VII.1985, 1360m, Larix forest; Prov. [0.89], 01 = 0.24-0.25 [0.243]. Kangwon, Kumgang-san Mts, above Kurjong water- Diagnosis ofworkers andquee«5.Workers: petiolar fall. No. 708-85, 18.VIL1985, 710 m, deciduous forest scale (seen in profile) relatively thick, not flattened at (all leg. M. Woyciechowski); 27 workers, North Korea, the top, approximately inversely U-shaped; when seen Onpho near Chogjin, No. 2355, 12.VIII.1959, leg. B. in front or from behind, it is only slightly narrowing Pisarski and Prdszyriski; 5 workers, Prov. Hamgyong- to the dorsal crest; head with convex sides, gradually J. pukto, distr. Orang, Changyon Lake, 17.VL1990, leg. and slightly narrowing anteriorly, and with distinctly R. Pisarska; 5 workers, Russia, viciiiity ofVladivostok, emarginate occipital margin; scape, mid and hind 5.V.1989, Quercus forest, leg. A. Radchenko; ibid., 2 tibiae not flattened, elliptical in cross-section; ratio of workers,1queen,1male,2.VI.1983,leg.A.Kupyanskaya; min/max diameters ofthe scape > 0.7; scape and legs 4workers, Russia, IsL Sakhalin, ChehovaMt,val. ofthe with decumbent pilosity only; promesonotal dorsum riv. Bureya, 29.VII.1986, leg. M. Nesterov; 4 workers, andoccipitalmarginwithrelativelyshortandabundant Russia, Primorky Region, Ussuriysky Natural Reserve, standinghairs. 10.VIII.1986, leg. A. Kotenko; 3 workers, Japan, Pref. Queens: petiolar scale (seen in profile) relatively Kanagawa, Doryosan, 10.Vn.l982, leg. M. Kubota; 5 thick, not flattened at the top, approximately inversely workers, "Japonia, Sauter" (IZK, MHZ, JUK. BPI). U-shaped;headwithconvexsides,graduallyandslightly Sinceall three castes of "oriental fuUginosus" were narrowing anteriorly, and with distinctly emarginate described and characterised comprehensively several occipital margin; scape, mid and hind tibiae not flat- timesbydifferentauthors(Wilson 1955;Yamauchi1978; tened, elliptical in cross-section; ratio of min/max Kupyanskaya 1989, 1990; Espadaler et al. 2001; Imai et diameters ofthe scape > 0.7; legs and scape with dense al. 2003), I do not provide formal description,just give decumbent pubescence only; head, alitrunk and gaster the diagnosis of workers and queens, and the draw- with abundant, but not very long standing hairs, and ings and measurements of type specimens. The main withwell-developeddecumbentpubescence. differential features ofit and I.fuUginosus are given in Distribution.RussianFarEast(Amursky,Khabarovsky Table 1. The differences between Lfuji and other East- and Primorsky Regions, Isl. Sakhalin, Southern Kurily AsianDendrolasius species are given inthe Key. Islands), north-eastern China, Korean Peninsula, Japan Measurements and indices of type specimens (the (allfourmainIslands);itisthemostcommonDendrolasius data of holotype are in brackets), [mean data are in speciesinthisarea. squarebrackets] workers: HLi= 1.19-1.43 (1.41) [1.33]; Etymology. The species is named after Fuji-san Mt., HL2 = 1.29-1.51 (1.51) [1.42], HWi = 1.18-1.43 (1.43) one ofthegreatest symbols ofJapan. [1.32], HW2 = 0.71-0.95 (0.92) [0.82], SL = 1.08-1.27 (1.27) [1.19], OL = 0.24-0.28 (0.28) [0.26], AL = 1.50- Key to Lasius {Dendrolasius) species of the East 1.68 (1.68) [1.57] mm; CI= 0.95-1.01 (1.01) [0.99], CLI Palaearctic = 1.06-1.10(1.07) [1.07],CWI= 1.53-1.57(1.55) [1.60], SIi = 0.86-0.92 (0.90) [0.89], SI2 = 0.88-0.93 (0.89) Though there are severalrecentversions ofthe Keys [0.90], 01 =0.18-0.21 (0.20) [0.19];queens: HLi= 1.36- to the identification of East Palaearctic Dendrolasius 1.40 [1.38]; HL2 = 1.44-1.50 [1.47], HWi = 1.40-1.46 species (YamauchiandHayashida 1968;Yamauchi 1978; [1.42], HW2 = 0.83-0.87 [0.84], SL == 1.26-1.27 [1.265], Kupyanskaya 1989, 1990; Imai et al. 2003), I proposed OL = 0.34-0.36 [0.345], AL = 1.90-2.04 [1.97] mm; CI somewhat modified version of the Key, including one = 1.03-1.04 [1.033], CLI = 1.06-1.07 [1.066], CWI = species more. Lfuji:;> tio. I LfuUginosus L^'r-.ll" workers workers - headusuallysomewhatlongerthanwide(CI0.95-1.01); - head lengthequaltoorlessthan itswidth (CI 1.00-1.03); - scaperelativelylonger(SI20.88-0.95); - scape relativelyshorter(Sb0.82-0.89); - standing hairsontheuppermargin ofpetiolarscalelonger,the - standing hairsontheuppermarginofpetiolarscaleshorter,the longesthairsdistinctlylongerthanthehalfofthemaximum longesthairsshorterthanthehalfofthemaximumdiameterof diameterofthescape; thescape; - decumbentpubescenceontheanterior(vertical)surfaceoffirst - decumbentpubescenceontheanterior(vertical)surfaceoffirst gastraltergiterelativelydense,distancebetween hairsdistinctly gastraltergiterelativelysparse,distancebetween hairsnotshorter shorterthanthehairs length; (usuallylonger)thanthehairs length; queens queens - eyeswith somewhatlongerhairs,lengthofthelongestones> - eyeswithsomewhatshorterhairs,lengthofthelongestones ^ mm mm 0.040 0.035 Table 1.Themaindifferentialfeatureso(LfujiandL.fuUginosus.