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Revision of the intertidal aleocharine genus Tarphiota (Coleoptera: Staphylinidae) PDF

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Preview Revision of the intertidal aleocharine genus Tarphiota (Coleoptera: Staphylinidae)

Vol. 107,No.4,September&October, 1996 177 EVISION OF THE INTERTIDAL CHARINE GENUS TARPHIOTA (COLEOPTERA: STAPHYLINIDAE)1 2 Kee-JeongAhn^ ABSTRACT: A systematic revision ofthe genus Tarphiota is presented. Tarphiota Casey is re- described.TarphiotafucicolaandT.geniculataareredescribed. Tarphiotapallidipessynonymized underT.fucicola. Lectotypeandparalectotypesaredesignated forT.pallidipes, T. debilicollis, T. hirsutula, T. iota, T. insolita, T. litorina, and T.seditiosafromCasey'ssyntypeseriesdepositedin theNMNH.AkeyisprovidedforseparationoftheknownspeciesofTarphiotaandillustrationsof diagnostic featuresarepresented. Members ofthe genus Tarphiota Casey are confined to the sandy beaches ofPacific North America. Tarphiota was first described and characterized by Casey in 1893. Maklin (1852) described Tachyusafucicola and Homalota geniculata from the coast of Alaska. Tachyusa fucicola was transferred to Homalota Mannerheim by Maklin in 1853. Casey (1893) described the new genus and species Tarphiota pallidipes from the coast of California. He be- lieved that the genus Tarphiota included H.fucicola and H. geniculata. Later, Casey also described six additional species from the Pacific coast ofAmerica. However, all sixspeciesweresynonymizedbyFenyesin 1920andT.pallidipes is synonymized under T.fucicola in this paper. Examination of the type series of T. fucicola (Maklin) and T. geniculata (Maklin) in the collections ofthe Finnish Museum ofNatural History allowed me torevisethis genus. In thispaper, Iredescribe Tarphiota Casey and thetwo valid species [T. fucicola (Maklin), T. geniculata (Maklin)], synonymize Tarphiota pallidipes Casey under Tarphiotafucicola (Maklin), and designate lectotypes andparalectotypesfromthesyntypeseriesofeachdescribedspecies NMNH. in the collections ofthe Tarphiota Casey TarphiotaCasey, 1893: 332, 1910:74;Fenyes, 1920:254;BernhauerandScheerpeltz, 1926: 596; Blackwelder,1952:374;Hatch, 1957: 145;MooreandLegner, 1975:489, 1976:535;Seevers, 1978: 132. DiagnosticCombination.Amongaleocharinegenerawith4-5-5 tarsal for- mula, membersofTarphiotaarerecognizedby thecombination of: integument 1 ReceivedFebruary26, 1996.AcceptedApril28, 1996. 2Contribution number 3149 from the Snow Entomological Museum (Natural History Museum, DivisionofEntomology),UniversityofKansas,Lawrence, KS66045, U. S.A. 3 SnowEntomologicalMuseum,Snow Hall,UniversityofKansas,Lawrence, KS66045. ENT.NEWS 107(4) 177-185,September&October, 1996 178 ENTOMOLOGICALNEWS ofhead, pronotum, and elytra granulose due to a fine-meshed, raised reticula- tion; antennomeres (Fig. 1) 8-10 transverse; head without infraorbital carina; mandible(Fig. 4)withbluntapex; labium(Fig. 6)withnarrowY-shapedligula; mentum with long v seta; separated mesocoxae; pointed mesosternal process; unmarginedmesocoxalcavities;anteriorandmiddletibiaewitharowofspines; elytraslightlysinuateapico-laterally,longerthanpronotum;hindwingspresent; and internal circular sclerites presentatbase ofmetasternum. Description.Small,length2.0-3.1 mm.Bodyshapenarrow,flattenedandparallel-sided.Body colorblack or dark brown. Microsculpture ofhead, pronotum, and abdominal segments faintly polygonal.Integumentofhead,pronotum,andelytragranuloseduetoafine-meshed,raisedreticu- lation,moreorlessuniformlypubescentwithshortmicrosetae. HEAD. Slightlydeflexed, =1.2timesaswideas long.Eyes withmicrosetaebetween facets, =0.5timesaslongastempora. Neckabsentbutbasal constrictionpresent. Infraorbital carinaab- sent. Microsetae relatively long, directed anteriorly, almost uniformly distributed. Integument granulose.Hexagonalmicrosculpturespresent.Temporashort.Submentumwithseveralsetaeand punctures.Antenna(Fig. 1)with 11 antennomeres,slightlyincrassatetowardstip;antennomeres2- 3eachshorterthanpreceding,4-7graduallydilated,8-10transverse. MOUTHPARTS. Labrum (Fig. 2)transverse, 9+9 majorsetae distinct, 3+3 additional setae present,sensillaepresent,puncturesdistributedanteriorly.Epipharynx(Fig.3)withabout6-8lat- eralporesoneachside,numeroustinypores,about6-8poresonapicalregion,about35poreson medianregionalongwithnumerousindistinctpores.Mandibles(Fig.4)asymmetrical,apexblunt and curved downward; right mandible with median tooth, absent from left mandible; serration absentfromapextomediantooth,manypuncturesscatteredventrallyanddorsally;prosthecawell developed, membranouswithfibrils. Maxilla(Fig. 5) withgaleaand laciniaelongate;galeawith innersurfacemembranous,apexwithlongfiliformsetae,outersurfacecorneous,apexwithnumer- ous delicate branching fibrils which havearow ofextremely delicate setae; laciniamoreorless acute, internal surface with comb composed ofasinglerow of9 well separated spines followed moreproximally byasmalldensepatchofsetae,and adistinctrowofnumerous setae,arowof poresondorsalsurface; maxillary palpus, robust,with4articles,article 3 incrassatedistally and longerthanarticle2,article4veryslender,narrowdistallywithfilamentoussensilla.Labialpalpi (Fig.6)with3articles,elongate,article3shorterthanarticle 1 andlongerthanarticle2,article3 incrassatedistally;ligulabifid,2tuberclesontip;apairofmedialsetae,twinpores,medianpores, distalpores,setalpores,realpores,basalporespresent,andabout30-35or5pseudoporespresent medially, 11-12or3 laterally. Prementumwithapairofcomb-likehypoglossae. Mentumalmost trapezoidal,slightlyemarginateanteriorly,threemainsetaepresentoneachside,vsetaerelatively long, severalsetaeandpuncturespresent. THORAX. Pronotum =1.2-1.3 times as wide as long, basal and apical lines about equal in length,sidesround;microsetaeuniformlydistributed;macrosetaeabsent;integumentgranulose,a rowofdistinctdarkpuncturesonbase. Patternofpubescencewithsetaesubparallel,thoseonthe anteriorhalfofpronotumdirected anteriorly and thoseon posteriorhalfdirectedposteriorly ina narrow median strip, lateral setae with anterior halfdirected antero-laterally and posterior half directed postero-laterally. Scutellum more or less octagonal. Hypomerafeebly inflexed, entirely visible in lateral aspect. Mesocoxal cavities narrowly separated; mesosternal process acute. Metasternumlongerthanwidthofmesocoxa.Tarsalformula4-5-5,internalcircularscleritespresent atbaseofmetasternum;anteriorandmiddletibiaewithdistinctrowofspines. ELYTRA. =1.2-1.3 times as long as pronotum; very slightly sinuate apicolaterally, longer laterally than medially, carina present on anterior medial edge ofelytra; microsetae numerous, uniformlydistributed,directedmoreorlessposteriorly;macrosetaeabsent;integumentgranulose. Hindwingspresent. ABDOMEN. General shape broad at base, sides uniformly converging to broadly rounded apex.TergitesIII-VIIIwithdistinctmacrosetae.TergitesIII-VImoderatelyimpressedatbase,ante- Vol. 107,No.4,September&October, 1996 179 riormarginoftergitesIII-VIdeeplyandbroadlyV-shaped.TergiteVIIIwith4or7pairsofmacrosetae. TergiteXwithsetanumerous,3-4majorsetaedistinct;posteriormarginnotwell sclerotized. AEDEAGUS. Medianlobe(Figs.7and 10). Paramere(Figs. 8and 11). SPERMATHECA. (Figs.9and 12). SECONDARYSEXUALCHARACTERISTICS. MaletergiteVIIIslightlysinuate. Typespecies. TarphiotapallidipesCasey [= T.fucicola(Maklin)]. Desig- nated by Casey (1910). Distribution. FromAlaska to California. Biology. Members ofthe genus Tarphiota inhabit the mid to upper littoral zone offine-grained sandy beaches, which is covered by only one or two high tides each month and contains decaying seaweed. Nothing is known oflarvae ordevelopment ofany species ofTarphiota. Remarks. Casey (1893) described a new genus Tarphiota for T. pallidipes and mentioned that members of Tarphiota resembled Alianta Thomson, but differed from members ofthis latter genus by having spinulose tibiae. Fenyes (1918) placed Tarphiota in the tribe Myrmedoniini (group Athetae) based on the 4-5-5 tarsal formula, antennae with 1 1 antennomeres, maxillary palpi with 4 articles, and labial palpi with 3 articles. The tribal name Myrmedoniini was later synonymized under the name ofAthetini by Seevers (1978). Bernhauer and Scheerpeltz (1926) classified the genus based on Casey's description and Fenyes' placement. Moore and Legner (1976) still placed Tarphiota in the Myrmedoniini(= Athetini).Seevers(1978)placedTarphiotainthetribeAthetini with Pontomalota Casey but noted that these intertidal genera were not neces- sarily related to each other and other members of Athetini. Ahn (in press) hypothesized that members of Tarphiota were closely related to members of Pontomalotaand ThinusaCasey andthatthesegeneramight formamonophyl- etic group within the tribeAthetini. Key to the species ofthe genus Tarphiota 1. Bodylength2.9-3.1 mm;abdominaltergiteVIIIwith7pairsofmacrosetae;spermathecawith narrowtube(Fig.9);paramereasinFig.8 T.fucicola Bodylength2.0-2.6mm;abdominaltergiteVIIIwith4pairsofmacrosetae;spermathecawith broadtube(Fig. 12);paramereasinFig. 11 T.geniculata Tarphiotafucicola (Maklin) TachyusafucicolaMaklin, 1852:306; Bland. 1865: 406. Homalotafucicola(Maklin), 1853: 182. Tarphiotafucicola Casey, 1893: 333; Bernhauerand Scheerpeltz, 1926: 596; Hatch, 1957: 145; MooreandLegner, 1975:489;LohseandSmetana, 1985: 286. TarphiotadebilicollixCasey, 1910:75.synonymybyFenyes 1920. TarphiotapallidipesCasey, 1893: 333;Fenyes, 1920: 254.NEWSYNONYM. Description.Length2.9-3.1 mm. Bodycolorblack,legsbrown. Head=1.2timesaswideaslong, integument granulose, microsculpture more ore less hexagonal. Labrum transverse. Epipharynx 180 ENTOMOLOGICALNEWS (Fig. 3)with=8 lateralporesoneachside. Labium(Fig. 6)with=12-13 medialpseudoporesand =11-12 lateral pseudopores. Pronotum =1.3 times as wide as long, integument granulose; sides round. Mesosternal process narrow, acute. Elytra =1.3 times as long as pronotum, integument granulose.Abdominaltergiteswithmicrosetaemoreorlessuniformlydistributed,severalmacrosetae present. Aedeagus. MedianlobeasinFig.7. ParamereasinFig. 8. Spermatheca.Tubenarrow(Fig.9). Type series. TachyusafucicolaMaklin,Lectotypedesignatedby Lohseand Smetanain 1985. Tarphiota debilicollis Casey: Lectotype, here designated, in the collection oftheNMNH,withlabelasfollows: "Caseybequest 1925;Lectotype, Tarphiota debilicollis Casey, Desig. K. J. Ahn, 1994". TarphiotapallidipesCasey: Lectotype,heredesignated, in thecollectionof the NMNH, with label as follows: "Casey bequest 1925; Lectotype, Tarphiota pallidipes Casey, Desig. K. J.Ahn, 1994". Paralectotypes, 8, same data as lec- totype. Material Examined. CANADA: British Columbia, Queen Charlotte Isl., LepasBay, Graham Island, 15July 1988,J.S.Ashe,exunderrottingseaweedonbeach(13, KSEM);Sidney(5,CAS); Victoria,Vane.,Hubbard&Schwarz(1,NMNH),Wickham(1,CAS).UNITEDSTATES:Alaska: GlacierBay, Unalaska,Van Dyke, 5 July 1907 (8, CAS); Nazan Bay,AtkaAleutian Is., 31 July 1907,V. Dyke(6,CAS), 1Aug. 1907(8,CAS).California: MarinCo.: PointReyes,NorthBeach, 15May 1991,K.J.Ahn&J.S.Ashe,exunderseaweed(1,KSEM);StinsonBeach, 14May 1991, K.J.Ahn&J.S.Ashe,exunderseaweedonbeach(17,KSEM);SanFranciscoCo.:SanFrancisco, 1 May 1910, F. E. Blaisdell (1, CAS), 27 May 1908 (1, CAS), 24 Nov. 1907 (1, CAS),24 Nov. 1918,E.P.VanDuzee(1,CAS);LongBeach,28Oct. 1916,J.O.Martin(4,CAS);SanMateoCo.: PacificBeach, 16May 1991, K.J.Ahn&J.S.Ashe,exunderseaweedonsandbeach(4,KSEM); Montara Beach, 16 May 1991, K. J. Ahn & J. S. Ashe, ex under seaweed on sand beach (32, KSEM); PomponioSt. Beach, 5 June 1991, K. J.Ahn,exunderseaweed(6, KSEM); Halfmoon BaySt.Beach, 16May,K.J.Ahn&J.S.Ashe,exunderseaweedonsandbeach(6,KSEM);Moss Beach, 12May 1910,F.E.Blaisdell(2,CAS),May 1910(1,CAS);SantaCruzCo.:Aptos, 19Nov. 1932,F.E.Blaisdell(2,CAS);HumboldtCo.:PrairieCreekRedwoodsSt.Park,GoldBluffBeach, 24July 1975,V.F.Lee(1,CAS);MontereyCo.:ZmudowskiSt.Beach,20May 1991,K.J.Ahn,ex underseaweed(3,KSEM);SalinasRiverSt.Beach,20May 1991,K.J.Ahn,exunderseaweed(43, KSEM);Carmel,8Sept. 1920,F.E.Blaisdell(4,CAS),24March 1919,E.P.VanDuzee(4,CAS); SanLuisObispoCo.:CayucosSt.Beach,22May 1991,K.J.Ahn,exunderseaweed(1,KSEM); SKa.nJt.aABhanr,beaxrauCnod.e:rSsaenatwaeBeadrb(2a,raK,SHEuMb)b;arSdan&MSicghuwealrIzsl(.,12,N0MJNunHe)1;91P0t.,SValWS.t.OBweeacnh,(23,JCuAnSe);19L9o1s, AngelesCo.:LosAngeles,April,Chittenden(1,NMNH),Aug.(2,NMNH),July(1,NMNH), 17 June 1937 (2, NMNH), 11 March 1934(1, NMNH); SantaMonica,March,A. Fenyes(2,CAS); Redondo, March, A. Fenyes (4, CAS), 4 June 1904, A. Fenyes (1, CAS); San Diego Co.: San Diego,20Aug. 1917,J.O. Martin (1,CAS); PortWatsonville,Cal,26Nov. 1932,F.E. Blaisdell (3, CAS); Cal, Hubbard & Schwarz (1, NMNH); Cal (4, NMNH); Del Monte, 7 June 1904, A.Fenyes(2,CAS). Distribution. FromAlaskatoCalifornia. Synonymy. I have examined Casey's syntypes of Tarphiota pallidipes. I could not find sufficient characters to separate Tarphiota pallidipes from T. Vol. 107,No.4,September&October, 1996 181 fucicola. Legcoloration,whichwaspreviously usedtodistinguish T.pallidipes from T.fucicola byCasey (1893), is notareliabledistinguishingcharacteristic. Tarphiotageniculata (Maklin) HomalotageniculataMaklin, 1852: 308;Bland, 1865:404. Tarphiotageniculata (Maklin), 1893: 334; Bernhauerand Scheerpeltz, 1926: 596; Hatch, 1957: 145;MooreandLegner, 1975:489;LohseandSmetana, 1985: 291. TarphiotahirsutulaCasey, 1910: 75.synonymybyFenyes 1920. TarphiotaiotaCasey, 1910:76. synonymybyFenyes 1920. TarphiotainsolitaCasey, 1910:76. synonymybyFenyes 1920. TarphiotalitorinaCasey, 1910:75. synonymybyFenyes 1920. TarphiotaseditiosaCasey, 1910: 76.synonymybyFenyes 1920. Description:Length2.0-2.6mm.Bodycolorentirelyblack.Head=1.2timesaswideaslong, integumentgranulose,microsculpturemoreorlesshexagonal.Labrumverytransverse.Epipharynx with=6lateralporesoneachside.Labiumwith=5medialpseudoporesand=3lateralpseudopores. Pronotum=1.2 timesas wideas long, integument granulose; apical line straight, basal line very slightly sinuate, sides round. Mesosternal process acute. Elytra=1.2times as long as pronotum, integument granulose. Abdominal tergites with microsetae more or less uniformly distributed, macrosetaenotdistinct. Aedeagus. MedianlobeasinFig. 10. ParamereasinFig. 11. Spermatheca.Tubebroad(Fig. 12). Type series. Homalota geniculata Maklin, Lectotype designated by Lohse and Smetana in 1985. Tarphiota hirsutula Casey: Lectotype, here designated, in the collection of the NMNH, with label as follows: "Cal, Casey bequest 1925; Lectotype, Tarphiota hirsutula Casey, Desig. K. J. Ahn, 1995". Tarphiota iota Casey: Lectotype, here designated, in the collection of the NMNH,withlabel asfollows: "Cal,Casey bequest 1925;Lectotype, Tarphiota iota Casey, Desig. K. J.Ahn, 1995". Paralectotype, 1, same data as lectotype. Tarphiota insolita Casey: Lectotype, here designated, in the collection of the NMNH, with label as follows: "Casey bequest 1925; Lectotype, Tarphiota insolitaCasey, Desig. K. J. Ahn, 1995". Tarphiota litorina Casey: Lectotype, here designated, in the collection of the NMNH, with label as follows: "Cal, Casey bequest 1925; Lectotype, Tarphiota litorinaCasey, Desig. K. J.Ahn, 1995". Paralectotypes, 2, samedata as lectotype. Tarphiota seditiosaCasey: Lectotype, here designated, in the collection of the NMNH, with label as follows: "Casey bequest 1925; Lectotype, Tarphiota seditiosa Casey, Desig. K J. Ahn, 1995". Paralectotype, 1, same data as lecto- type. Material Examined. CANADA: British Columbia, Queen Charlotte Isl., Lepas Bay, Graham Island, 15July 1988,J.S.Ashe,exunderrottingseaweedonbeach(35, KSEM);Massett,nodate. Rev.Keen(2,CAS);Sidney,nodate,A.Fenyes(1,CAS). UNITEDSTATES:Alaska: NazanBay, 182 ENTOMOLOGICALNEWS 6 Figs. 1-6. T.fucicola(Miiklin). 1,Antenna,dorsalaspect;2, labrum,dorsalaspect; 3,epipharynx, dorsalaspect;4,rightmandible,ventralaspect;5,maxilla,dorsalaspect.6,labium,dorsalaspect. Scale,0.1 mm. Vol. 107, No.4,September&October, 1996 183 9 10 12 Figs. 7-12. T.fucicola (Maklin), 7-9 and T. geniculata (Maklin), 10-12. 7, Median lobe, lateral aspect; 8, paramere, lateral aspect;9, spermatheca,dorsalaspect; 10, median lobe, lateralaspect; 11,paramere,lateralaspect; 12,spermatheca,dorsalaspect. Scale,0.1 mm. 184 ENTOMOLOGICALNEWS AtkaAleutianIsl., 1 Aug. 1907,VanDyke(3,CAS).Oregon:LincolnCo.: 5.0mi. S.Waldport,26 Aug. 1986,J. S.Ashe,exrunningonwavesweptshore(5,KSEM);4.0mi. N.Yachtshwy 101,27 Aug. 1986, J. S. Ashe, ex on sand beach (39, KSEM). California: Humboldt Co.: Prairie Creek RedwoodSt. Park,GoldBluffBeach,24July 1975,V.F.Lee(6,CAS);MarinCo.: BolinasPoint, 1.6mi.duewestofBolinas, 16June 1974,V.F.Lee(8,CAS),20April 1975(7,CAS),9June 1974 (2,CAS);BolinasPoint,2June 1974,V.F.Lee(4,CAS);StinsonBeach, 14May 1991,K.J.Ahn & J. S. Ashe, ex underseaweed on sand beach (2, KSEM); Point Reyes, North Beach, 14 May 1991, K. J.Ahn &J. S.Ashe,ex underseaweed (1, KSEM); PointReyes,Nat. SeashoreWildcat Beach, 1 June 1975,V.F.Lee(2,CAS);SanFranciscoCo.: SanFrancisco,nodate,F.E.Blaisdell (4, NMNH),Aug. (16, CAS),July (2, CAS), 9 Sept. 1909 (1, CAS), 27 May 1908 (1, CAS), 24 Nov. 1907(4,CAS),May(1,CAS),6Aug.,Wickham(4,CAS), 1 Aug.(2,CAS);SanMateoCo.: Halfmoon Bay St. Beach, 16 May 1991, K. J. Ahn &J. S.Ashe, ex underseaweed (10, KSEM); MontaraBeach, 16May 1991,K.J.Ahn&J.S.Ashe,exunderseaweedonsandbeach(1,KSEM); PomponioSt.Beach,5June 1991,K.J.Ahn,exunderseaweed(12,KSEM);MontereyCo.:Asilomar St. Beach, 20 May 1991, K. J. Ahn, ex underseaweed (10, KSEM);AsilomarBeach, 31 March 1968,J.B.Evans,exunderwetseaweed(1,KSEM);SalinasRiverSt.Beach,20May 1991,K.J. Ahn, ex underseaweed; Zmudowski St. Beach, 20 May 1991, K. J.Ahn, ex underseaweed (41, KSEM);Carmel,24March 1919,E.P.VanDuzee(1,CAS),8Sept. 1920,F.E.Blaisdell(8,CAS); SanLuisObispoCo.: PismoSt.Beach,2June 1991,K.J.Ahn,exunderseaweedonsandbeach(4, KSEM);CayucosSt.Beach,22May 1991,K.J.Ahn,exunderseaweed(23,KSEM);MorroStrand St. Beach,22May 1991, K.J.Ahn,exunderseaweed(8, KSEM);SantaBarbaraCo.: SantaBar- bara, 16June 1931,F. E. Blaisdell (1, CAS); Pt. Sal St. Beach, 3June 1991, K. J.Ahn,ex under seaweed(42,KSEM);CarpinteriaSt.Beach,23May 1991,K.J.Ahn,exunderseaweed(4,KSEM); Figs. 13-14. 13, Tarphiotafucicola (Maklin), habitus. Length: 3.0mm; 14, Tarphiotageniculata (Maklin),habitus.Length:2.4mm. Vol. 107, No.4,September&October, 1996 185 SanMiguelIsl.,20June 1910,V.W.Owen(2,CAS);VenturaCo.:SanBuenaventuraSt.Beach,23 May 1991,K.J.Ahn,exunderseaweed(5, KSEM);LosAngelesCo.: LongBeach,28Oct. 1916, J. O. Martin (3,CAS); Redondo, March,A. Fenyes (3, CAS),April (1, CAS), July (1, CAS); S. Pedro, Aug. (2, KSEM): Cal (3, NMNH); San Diego Co.: San Diego, Hubbard & Schwarz (1, NMNH),20Aug. 1917,J.O.Martin(8,CAS),22Nov. 1890,F.E.Blaisdell(5,CAS), 19Jan. 1928 (4,CAS),nodate(2,CAS),nodate,Wickham(1,CAS);SilverStrandSt.Beach,28May 1991,K. J.Ahn,exunderseaweed(8, KSEM);Del Monte,June,A.Fenyes(2,CAS);Caseybequest 1925 (6,NMNH). Distribution. FromAlaska to California. ACKNOWLEDGMENTS Ithankthefollowingpeoplefortheircontributionstothisproject. D. Kavanaugh(California Academy ofSciences; CAS), J. Muona (Finnish Museum ofNatural History; FMNHC), G. N. House (National Museum of Natural History; NMNH), and J. S. Ashe (Snow Entomological Museum, UniversityofKansas; KSEM)whogenerouslyprovidedmewiththeopportunitytoex- aminethecollectionsincludingthetypeseriesofspeciesofTarphiotabyarrangingfortheloanof specimens. IalsothankJamesS.AsheandSteveW.Lingafelterforreadingandprovidinghelpful suggestions on the manuscript. This research was supported by Snow Entomological Museum DevelopmentFund,Univ.ofKansasGeneralResearchFundAward#91-162,andNSFGrantDEB- 9521755awardedtoJamesS.Ashe. LITERATURECITED Ahn, K. -J. in press. Revision and systematic position of the intertidal genus Thmusa Casey (Coleoptera: Staphylinidae:Aleocharinae). Ent. Scand. Bernhauer, M. and O. Scheerpeltz. 1926. Coleopterorum catalogus. Pars 82, Staphylinidae 6:499-988. Blackwelder,R.E. 1952.Thegenericnamesofthebeetlefamily Staphylinidaewithanessayon genotype. Bull.U.S.Natl.Mus.200:1-483. Bland, J. H. B. 1865. Compiled descriptions ofNorthAmerican Staphylinidae. Proc. Entomol. Soc. Phila.4: 391-425. Casey,T.L. 1893.ColeopterologicalnoticesV.Ann.N.Y.Acad. Sci.7:281-606. Casey,T.L. 1910.NewspeciesofthestaphylmidtribeMyrmedoniini. Mem. Coleop. 1: 1-183. Fenyes,A. 1918-21. Genera Insectorum, Coleoptera, fam. Staphylinidae, subfam. Aleocharinae. fasc. 173a,b,c: 1-453. Hatch,M. 1957.ThebeetlesofthePacificNorthwestPart II: Staphyliniformia. Univ.Wash. Pub. Biol. 16:1-384. Lohse, G. A. and Smetana. 1985. Revision ofthe types ofspecies ofOxypodini and Athetini (sensu Seevers) described by Mannerheim and Maklin from North America (Coleoptera: Staphylinidae).Coleopt. Bull. 39(3):281-300. Maklin, F.G. 1852. Descriptionofnewtaxa. In: Mannerheim, C. ZweiterNachtrag zur Kaefer- faunaderNord-AmerikanischenLaenderdesRussischenReiches.Bull.Soc.Imp.Nat.Moscou, 25:283-387. Maklin,F.G. 1853.Descriptionofnewtaxa.In:Mannerheim,C.DritterNachtragderAleutischen Ins. Bull. Soc. Imp.Nat. Moscou,26:95-269. Moore,I.andE.F.Legner. 1975.ACatalogueoftheStaphylinidaeofAmericaNorthofMexico (Coleoptera). Univ.Calif. Div.Agric.Sci.Spec. Publ.3015. Moore, I. and E. F. Legner. 1976. Intertidal rove beetles (Coleoptera: Staphylinidae). In Cheng [ed.]. Marineinsects. NorthHolland Publishers,Amsterdam. Seevers,C.H. 1978.AgenericandtribalrevisionoftheNorthAmericanAleochannae(Coleoptera: Staphylinidae).FieldianaZool. 71: 1-289.

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