Vol. 6, No. 3-4, September - December, 1992 151

A revision of the New World species of Norrbomia (Diptera: Sphaeroceridae), inch.ding all American species previo'.sly placed in Borborillus

Stepllell A. Malsllall Department Of Environmental Biology, University of Guelph Guelph, Ontario, Canada, N1 G 2W1

and

Allen L. Norrbom Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA, c/o National Museum of Natural History, NHB 168, Washington, D.C. 20560

Abstract The New world species formerly placed in Bilrb67ilbts I hlda CspbaA1'Oi'Aridae' I :Opmmyzinae) are trans- ferredtoNorrbomiaPapp. Theseare:N. lacteipennis(Malloch), n.comb.,N. jitmipennis(Stenhammar),n.comb., N. {tigipennis (Spuler), n. comb., N. scripta (Malloch), n. comb., and N. sordida (Zetterstedt). Norrbomia jitlvipennis, N. singusta, N. mexicana, N. triglabro, and N. yukonensis are described as new. Borboros articus MallochissyrwnymizedwithNo {amipennis, andBot burassingalaris Spaler is syrwnymized with No scripta. All tenNewWorld speciesofNorrbomia arekeyed. illustratedanddescribed. Theirrelationshipsarediscussed and a cladograrn is provided. Species in two ofthedefined clades arekleptoparasiticon dWIg rolling scarab beetles.

Introduction Materials and methods Most Holarctic species of Sphaeroceridae with a Morphological terminology follows that of closed basal cubital cell, an apiCO"v'entral spur on the MeAlpine (1981). As in most CopJ:6mYzinae, the hindtibia, a singlerowofpostocularbristles.andonly pattern of microtrichia on the frons, gena, and tho- fourscutellarbristlesuntllnowhavebeenincludedin racicpleuronvarlesconsiderablyamongthespeciesof the genus Rorborilb1.s Duda Papp (1988) recently Norrhomia. and isuseful todistinguish them (Figs 4- discovered that the type speCIes of BorbOnllUS, the 25). ThischaracterlSmoredifficUlttousemN01TbOmw European species B. uncinatus (Duda), as well as B. than. in most other oopromyzine genera, hOVlre>.rer, vitripennis (Meigen), are not closely related to most becauseinsomespecies(e.g.N.lacteipennis(Malloch), other IIoiarctic speciesplacedinBorborillus, butale Figs. 1 2) the cuticle is more striated or rugose than insteadrelatedto a primarilyAfrotropical groupthat normal, andthe bareareasarethereforenotasshiny mcludes MetabOrbOros Vanschuytbroeck, Gymno­ anddonotcontrastaswellWIth thell1lcrotrichose (or metopinaHedicke,DudaiaHedicke,and AfroborbonlS pruinose) areas In some species, the exact size and Curran. The name Borborillus therefore cannot be shape of the bare areas are difficUlt to see with the usedfortheNorthAmericanspeciespreviouslyplaeed light miooscope and were con.'irmed only by using inthatgenus. Thesespecies,inadditiontomostofthe scanning electron microscopy. Old World speCIes preVIously m Borborillus, should Wnen discusSIng the genal bare area we are now be treated within the geWIS Noribomia Papp, referring mainly to the anterior part that extends proposed for N. mdun Papp and a number of alongthe ventralmargIn ofthe eye. There IS always Afrotropieal species revised by Papp (1988). a vertieal bare area along the posterior edge of the gena from the ventral margintobehindthe eye. The exL:reme ventral margin also usually has a separate 152 Insecta Mundi

Figwee 1-2. NOrrOOmia l=teipermi8. 1, high magnification ofocellar region; 2, border of microtrichose and bare areas on ~epi8temum(shows rugose areaofcuticleandwhybordersofmicrotrichoeeareasaredifficultto 110O insomespecies). very 1l81TOW bare area. The anteriorpartofthe bare Genus Norrbomia Papp area on the frons lateral to the ocelli, if present, is often difficult to see because of cuticular striations Norrbomin Papp 1988:394. Type species, N. iru:iim (Fig. 1). This bare area therefore often appears to be Papp, by original designation. smaller than it really is. We refer to the slender (inpart)Copromy:w. (BorborillU8): Richards 1965:7~, eclerotizedbasalpartofthespermathecathetextends 1967:2. to the slightly expanded, membranous spermathecal (in part)Borborillu8: authors. duet as the spermathecal neck. The spermathecae also have cylindrical basal or apical apodemes. Genericdiagnosis. Relativelyamall Copromyzinae In the Material examined sections, specimens (00. 2-4 rom long). Head: Face deeply concave, often listedwithoutmuseumacronymsaredepositedinthe with pale, membranous medial strip. Postocular Universityof Guelph collection. Acronyms for other bristles insingle row. Tlwm:c: Scutellum short, with institutions are as follows: American Museum of 4margina1bristles only. Fore first taraomereofmale Natural History (AMNH); BMNH - Natural History withstout, dark, curved, apicovent>-al spur. Midtibia Museum, London (formerly British Museum (Natu­ with anterovent>-al bristle below middle and ring of ral History»; CAS - California Academy ofSciences; preapical and apical bristles. Hindtibiawith curved, CNC - Canadian National Collection; FMNH - Field ventrally directed, apicoventral spur, thin preapical Museum of Natural History; FSCA - Florida State dorsalbristleandanterovent>-al bristlebelowmiddle. CollectionofArthropods;GUE- UniversityofGuelph; Maleabdomen: Hypandriumwithlong apodeme and lNHS - Illinois Natural History Survey; MCZ - Mu­ well developed arms; synsternite6+7withoutmedial seum of Comparative Zoology, Harvard University posterior lobe; postphallic sclerite (ec1erite between (MCZ);NRS-NatumistoriskaRiksmuseet,Stoekholm; subepandrial eclerite ('sternite 10") andbaaiphal1us) OhSU- OhioStateUniversity;UKaL-SnowEntomo­ absent; basiphal1us with well developed epiphallus; logical Museum, University ofKansas; USNM -Na­ epandrium with lateral clefts; cercus long, distally tional Museum of Natural History, Smithsonian In­ modified andconnectedtoepandrium; surstyluswith stitution; WSU - Washington State University; ZlL­ flattened inner lobe. Female abdomen: 2 spermathe­ Zoological Institute, Lund; ZMHU - Zoologisches cae,eachwithslendereclerotizedneckatleastaslong MuseumanderHumbolt-Universitat. Becauseofthe as main partofspermatheca (except in N. hispanica largenumberofspecimensexaminedofN. frigipennis (Duds» and with large basal or apical apodeme. andN. sordida, wehavenotlistedcompletecollection data for these species, Vol. 6, No. 3-4, September - December, 1992 153

Table 1. List of characters and states used in phylogenetic anal}sis of ~"lorrbomi4. State 0 iB hypothesized as plesiomorphic.

1. Basiphallus .0)slender, lengthusuallymarethan3.5xwidth(in 17. Hypoproct . 0) anterior margin not strongly concave, without latemlvie",..); 1) shOl't and Bteut, lengthuBUally less than 3.0. seplU'llte selerite; 1) anteriCll' margin strongly eE>neave, with (++) separate bIfid sclente. (+++) 2.Malecercus•0)simpleandsmall; 1)modified,usuallylarge.(+++) 18. Epandrium setation • 0) dOl'8al and lateral setae similar; 1) 3. SpennaLheea neck • 0) absent or shorl:et than main part of dorsal setae less than halfaslollg as moBL lateral seLae. (++) spennatheea; 1) &8 long &8 main part ofspennatheea. (++) 19. Base of male _s and bel'deFing pal't of epandrium 0) 4. SUl'IItylus • 0) WIthout bIadelike mes81 lobe; I) WIth bladelike WIthoutclust.erofbristles; I)WIthclust.erofnumerousbristles. mesal lobe. (+++) (++) 5. DistiphaHus 0) without basal sclerite, 1) with basal sclerite. 20. Tarsi color • 0) similm to Libiae, 1) paler th8nLibiae. (+) (+++) 21. Female sternite 4 • 0) entire; 1) bipartite; 2) tripartite; 3) 6 IliBtipha 1I11s basaI sclerlt.e • 0) diVidAdj 1)1IndiVidAd (+++) cp.adrlparlit.e or aeoondariIy bipartite Stat.es 1, 2 and 3 are 7. Spennatheca basal apodeme • 0) smallorabsent; 1) large. (+++) hypothesized as independently derived from state O. (++) 8. Frena miaettiehialpattern 0)withbare spats borderingoeelli; 22. Female tergites 6 anti 7 0) entire to mOOer ately trilobed 1) entirelY microtricli08e; 2) entirely bare. States 0 ana 2 are posteriorly(incisions extendedless than213 distance toante· assumed to be mdependentty derIved from state O. (+) nor margm); 1) very deeplY lobate postenorlV ImClSIOll8 9. Hind tibia· 0) with stout anteroapical bristle; 1) without stout extended mare than 2/.3 distance to anterior margin). (++) antlll'Qapical bristle (+) 23 Female teIlJite 6 0) not bipartite; 1) bipartite. (+) 10. EPlproct setation . 0) more than 1 pall'; 1) I paIr. (+) 24. EPlproct • 0) not elongate, not bllobed anteriorly; 1) elongate, 11. Distiphallus basal sclerite • 0) lateral parts not sheathlike; 1) bilobed anteriorly. (+) lateml parts aheathlike. (I I I) 25. IIypoproct 0) anteriw edgenotbilobed; anteromedialpartnot 12. Spennatheca shape· 0) spheroid; 1)teardrop shaped (+++) ooncave; 13. F'em81e sterriites 6 and 'I . 0) anterIorly WIthout transverse 1) antenor edge bdOlJed, anteromediBl plll"t concave. (+) groove; 1) anteriorly with transverse groove. (+++) 26. KIeptoparasitic on 8C8rBb beetles· 0) no; 1) yes. (++) 14. Suratylus,poateloventralpart O)withoutzow ofatoutbriatles, 27. Bristles ofpoateroventralpartofauratylua· 0) allOt L1;0 m0der­ 1)with distinctrow oflong, stout, inwlll'dly directed bristles. ately long; 1) extremely long. (++) (+++) 2B Male cereus. 0) single lobed arnot with jnner lobE' longer than 15. Anepiaternum bare area • 0) BmBlI to moderate sized, dOl'8al outer lobe; 1) bilobed or notched, with long inner and shorL marginmore ax less Ie.elwithor .entra! to •entralmarginof outer lobe. (++ I) spiracle; 1) large, d01'88l margin levelwith ordOl'88.1 to dOl'8BI 29. Pleura of female segments 6 and 7 • 0) simple; 1) rugose or marginofapiracle;2)entirelybare.StatesOand2are 88Sumed striate. (++) to be independently derived from state O. (+) 30. Wing • 0) not white; 1) white. (+) HI. Male _s 0) Bot mised poateriCll'1y; 1) miaed posteriorly, with kriOb ornage. (++)

Biology. ~\To"bomia species are probably all eop associated with scarabs (see Comments undet that rophuguus. SOUle species, such as tire cosmopolitan, lM'CiOO. synanthropic N. sordida (Zetterstedt), appear to be associatedWith unglllate d1mglmder a Wide range Of RelationshiPs. we present here an analySis Of the conditions, yet are not as abundant as some relationshipsofNorrbomiatoothergeneraofthetribe Limosininae (8Phaeroooridae) assoeiated with simi GOPromyzini. Althoughwehavenotstudiedalloft.he lar habitats. Norrbomia {umipennis (Stenhammar) OldWorldspecies, we discusssometentativehypoth- and N. singusta, n. sp., have been collected from esesofrelationshipsamongtire speciesofNo17 bomia, mammal bwrows, and the fOlmer bred from them. with emphasis on the New World fauna. Norrbomiayukonensis, n. sp.,probablyalsooccursin Characters are listed in Table 1, and the mammal bUrrows Perhapsthemostiliterestiiigana distributiOns of the CharaCter states are mown in best known ofthe Norrbqmia species are those that Tables 2 and 3. The matrix inTable 2 was analyzed arekleptoparasitieonthedungeaehesofSearabaeidae using Hennig86 (Farris 1988). Resulting bees are (Coleoptera). Membersofthe lacteipennisgrouphave shown in Fig. 3, with homoplasies indicated by (.), long beennoted as riding on adult scarabs (Moulton and reversals by (-). Two runs were made using the 1880, Lab 1915, SteySkal 19'11) SiilinSld (1983, imPlicit enumera.·tiOnoPtiOn (ie*). Thefirst analysis, 1984) described the behavior and life history of two in which the characters were equally weighted, membersofthisgroup,N. {rigipennis(Spuler)andN. produced two equally parsimonious trees 26 stepsin (ulvipennis,n.sp.(thelatterasBorborillussingularis). length (Fig. 3A·B). In the second analysis, each iVorrbomia bVI ipta (Malloch) also has been found character was assigned a weight of 1, 2 or 3, as indicated bythe numberofpluses jn Table 1 and Fig 154 Insecta Mundi

Table 20 Character statedisLt manansinspeciesgroups of Table 9. Character state disLt mutions in New 'Norid Norrbomia and inrelated genera. Numbers ofcharacters species of the scripta, costalis and frigipennis species and sLates refet to Table 1. gIOUpS of ...\'oIT'bomia. Numbers ofeharaeters and states refer to Table l. Character numbers _. . Taxa '63 4 56 7 a 901 63 4 5 67 B 9 0 Taxa 21 22 23 24 25 26 27 28 29 30 cr ~a o 0 0 0 0 ? 0010000000100 000 -u -u -r 'J 'J -u -u v v v Richardsia 1000000 0 010000 0 o 0 0 0 0 11nnnnn? n1nnnn? nnnnn mexicana 1 0 0 0 0 ? 0 0 0 0

.J.nD.J.C8 gr. LLLLLUU L LULLUU U UUUUU -"- v v -"- ~ =stalis gr. 1111111 1 100000 0 1 1 0 0 0 fumipennis 3 0 0 0 0 0 0 0 0 0 qrdnt-;o rrr 1 1 1 1 1 n n 1 n n n --::;- --;:: --;:: --;:: --;:: --;:: --;:: n n n frig:ipennis gr1 1 1 1 1 1 1 0 010000 0 o 0 1 1 1 triglabra ??? ? ? ? 0 0 ? 0 ,~~.. :_~--"- n 1 1 1 1 1 1 1 n 1 ~ ~ ...... 1 1 1 3. Weights were assigned accordingto the systemof fulvipennis 2 1 1 1 1 1 1 1 1 0 I MIl'> C'.onea ..,.. $In:! liD' weig,ht usually uniq,ue characters',secoded()tho + are low weight attrtbutes fOr Which pOlarity or homplogy are more likely to be misinterpreted, or Papp(1988)proposedthegenusNorrbomiaforN. which are commonly subject to homoplasy in other indica Papp and the P8.IeotropicaI species that were genera ofSphaeroceridae. Trees BO-D were the only previouslyplaeedinBorborilktsDuda. Henotedthat trees that resulted from the second analysis. Using Borborillus, as previously composed, was polyphyl­ unweighted characters, trees 3C-D are 26 steps in etic. The type species, B. uncinatus (nuda); and B. length, one step longer than trees 3A-B. But using vitripennis(Meigen)areparlofa monophyleticgroup weighted GharaGters, trees ac D are 51 steps lOng, aIsoi1icl.udillgDudaia,AtrobOrlJOros, CJYmnometopina, whereastrees 3A-B areeach54stepslong. InFig. 3D and Metaborboros (Norrbom and Marshall 1988). itis equally parsimonious for character 10, state 1 to Papp (1988) suggested that most other species for­ be an independent apomorphy in Richardsia, in merly placed in Borborillus probably also belong in AchaEtothomx,andinthefrigipennis +scriptagroups, Norrbomia. OurstudyconfirmsthisforalloftheNew rather than as shOWn on the tree, but thiS dOes not World species Trile BorbOrilhLS species are nOt change the tree topology or length. known from the Americas. Inbothoftheabo-."e analyses,weeodedCrumomyia The relationships of Norrbomia species to other as having state 0 for characters 8 and: 15, although Copmmyzini have not been discussed previousl), states 0 and 1 ofboth characters occurinthe genus. other than the statement by Papp (1988) that they We oonSider the states ended 0 man:! likely to be Jack apomorphies ofthe male genitalia found in the plesiomorphic for Norrbomia because they occur in Borborillus -Metaborboros clade. Infact, Norrbomia Riehard8ia. Also, Alloborbo1U8 Duda, wbiOO is the 8.Iso lackS apomorphies ofthe largergroup composed sistergroup ofCromomyia, has state0 ofcharacter8 of this clade plus Lotophila Lioy and Copromyza and an intermediate condition between sLates 0-1 of Falh~n (NcnTbom and Kim 1985). character 15 Ifthe polarityoroneorboth characters Norrbomiaappearstoform amonophylettcgroup is reversed by changing the state for Cromomyia, with AchaEtothomx Hedicke (Fig. 3A-D). This hy­ trees Withthe same:relationshiPs as Fig. 3C D result potheSis Of relationshiP is suppoi'ted primarilY bY ifthe characters are weighted. Ifthe characters are their modified male cercus (character 2), which is unweighted, various bees result, including some usuallyelongate, with lobes or depressions such: that with RichardsiaratherthanAchaEtothomxas sister itsshapeisnormallyspeciesspecific. Thisdiffersfrom group ofNorrbomia. the plesiomorphic state found in most Copromyzini, in which the cereusissma]) and simple Somespecies

Figure 3. Relationships of 6"lo1"1'OOmia. A D, among species groups of Av,1"1'OOmia and related genera; A B, based on unweighted characters; C·D, based on weighted characters. E, relationships among species of frigipennis group. F, relationships among species of scripta group. Numbers refer to characters listed in: Table 1. -~" symbols represent character weights, -." symbols represent homoplasies, -." symbols represent reversals. Vol. 6, No. 3-4, September - December, 1992 155

======::::'::'-(~1++'~10+)J[(2+++)i ~(8 2+.15 2+) =X ~ ======~('3+~+.'4+++"5+)~ scripta gr. 18++ 19++ 20+ r. (16++.)==== costalis gr.

A 10- indica gr.

,,'

~ ~ ~ ~ ~.~ '''' ~ , ...... v ...... =12...... +)= rr==113+++ 14+++ 15+)= scriota or. r . ...II 7*** ~(18++.19++.20+)=== frigipennis gr.

4+++. -r===(6***.7***.)==== costalis gr. .,., 0 '" ..n \ 1C .1'7 10- k (11+++ 12+++ = indica g r. Richardsia

'(l++.lO+)= ,l:S.:l+.1~.:l+ Acnaecocnorax

.~ ~ 1~ 1 1 -'I.' ~ • J ...... ",. 12+++ =\,j++.,= =\.L,j+++• .1t,l+++ • .1:>+) scrl.pca gr. 4+++. C 0 1n "\n \ ./:. '" .. . '" 7+++ .~ ':1 - ':>s a .s 16++.17+++ Rl.chardsl.a

(1++. )= (8.2+.15.2+) Achaetothorax 1n ==(:l+++)= ,11+++. 1~+++ l.ndl.ca gr.

•J .... "'~ . 4+++ • II ..,. = h 1=1 . ~ ... nr 8* 9* 7+++ 16(8- )==11 14+++.15+ ...... "'~ . - 10+ 20+ triglabra

= (30") l.actel.pennl.s = =(22++[/3*. )J I(30*)= frigipennis =(21.1++)=Jl ------...-.§~~~~,.---c.~...... ,.;-';-;-:+------.E(21.291 ~) (62 **.) scri ta ) fulVl.pennl.S F 30* E 28+++ 2++ 156 Insecta Mundi ofMetaborborusandGymrwmetopinaalsohavemodi­ though a dorsally projected sclerite is present just fled male cerci, buttheirshapes are different and we apical to this do notcoIlSld.erthis a synapomorphyforthosespec1es The NorrbOmta speCIes we exanuned appear to and Norrbomia and Achaetothorax. The relatively form four clades, here tenned the indica, 008talis, short basiphallus (character 1) is another probable scripta, and frigipennis groups. The relationships synapomorphyforNorrbomiaandAchaetothoraxand among Urese species groups are uol; well resolved. possibly also Richardsia Papp, a monotypic genus Characters 8, 9 and 10 suggest that the indica and whoserelationshipsarepoorlyunderstood. Theratio costalis groups are SIster taxa ()'ig. 3A-B), whereas of hasiphaJ1us length tn nalTOwest width in lateral characters 6 and 7 indicate that the indica gEe'lp is VIew IS usually less than 2.0 In Rtchardsta, the SIster taXon of a clade mCluding the other th1'ee Achaetotlwrax, and within Norrbomia, inthe indica groups(Fig. 3C D). Thelatterhypoihesisisprefen.ed group, the scripta and frigipennis groups, and some when characterweights are considered. Characters speciesofthe costalis group {t,.g. gravis (Adams». In 6811\17, which are complex8ud unigDe 1;0 Ute costalis. otherElpecies ofthe costalis group, suchas N. rostalis scriptaandfrigipennisgroups,arelesslikelytobetb.e (Zetterstedt), N. jUmipennis (Stenhammar), and N. resultofhomoplasythancharacters SOLO. Character yukonensis, n sp., itis 2.5-3.0, and as much. as 3.5 in 8, the miGrotrichial pattern of the frons, is highly N. mtidifrOns (DUda). In most otherCopromyzinae, variable withinthe Copromyzinae. Forexample, the theratiois gre~thana.5, andusuallygreaterthan size and presence of the bare areas vary within the 4.0. The basiphallus is short (ratio less than 3.5) in genera Crumomyia Macquart, Archiborbo1U8 Duda, some species ot Gymnometopina, Dudaia, and Lotophila, and Metaooroo1U8. We tentatively con­ Metaborborus, but its shape is different and other siderststeo(fronswith2-3smallbarespotsbordering character state distributions mdicate that this IS the ocelli) to be pleSIomorphic beCause thiS state homoplasy. The reduced number of setae on the ooours in Richardsia and is the more eommon state epiproct (Character 10) might be another within the Copromyzini. Both characters 9 and 10 synapomorphy for N017'oomia + Achaetothmux + involve Ute lossofbrisUes. Characterlossisgenerally Richardsia. However,thereisconsiderablehomoplasy considered more likely to occur through homoplasy inthischaracler(thepleSIomorphicstateIS presentm thanthemdependentdevelopmentofSilriilarcharac­ the indica and costalis groups of Norrbomia), and ters, so these charaeters are assigned low weight. because itis a loss characterwe have assigned it low Although the weighted character trees suggest weight in the analysis. thatthe indicagroup is the siStel group of Ute resl; of The monophyly of Achaetothorax is well estab­ Norrbomia,relationships amongthescripta, costalis, lishedbyasui1eofsynapomnrphies(l'appandNorrhorn andfrigipennl'.Rgronpsarennresolved Thereisweak ] 992) The monopbyly of Norrbomia is indicated by evidence that the scripta and frigipennis gEOUpS are three synapomorphies (clUlriiCters 3-5). The sper­ themoreClosely relatedofthe th1'ee groups (Fig. 3D). matheca has a very slender, selerotizedneckat least If charaeters 8 and 9 and character 10, state 0 are aslongasthemainpartofthespermatheca(character assumedto besynapomorphiesforNorrbomia,rever­ 3). Tireneckis very shortinl{. h@W'tiooofLhe indioo sal of these characters are aynapomorphies tor the group, whichwe suggestis due to reversal. Itis also scripta + frigipennis groups. Because these are low sometimes shorter than the main part Of the spar- weight Characters and a triChOtomy Of the thnle matheca in N. (ulvipennis. Some species of groups is equally parsimonious (Fig. 3C), we do not Achaetothorax, Gymnometopina, and Metaborbo1U8 consider this evidence very significant. have an elongate neck, bul; it; usuaRy is uol; longer The indica group is primarily PaleoLropical; N. than the main part ofthe spermatheca, and itis not 80rdidaistheonlyspeciesofthisgrouppresentinthe asslenderasin Nott1Jnmia,wbich snggeAtsthisisdlle Americas,piObablythroughspread6yman Bypath- to convergence. The othertwo synapomorphies are: esized synapomorphies for this group include: the thesurstylushasa blade-likemesaIlobe(Cliaraeter4), baSal sclerite of the distiph8Ilus with sheath-like astroetureuniquetoNorrbomia; and,thedistiphallus lateral parts, the medial part bilobed 01 tJ: uncate hasa large,dorsallyprojeeted,uniquelyshaped,basal (character11); andthe spermathecateardropshaped sclerite (character5). The lattermay be homologous (cha.ract.erl2). Manyofthespeciesoftheindicagroup with similarstructuresinothercopromyzinegenera, are densely microtrichose and have a genal pattern bUt its Shape is unique to NOrrbOmia. OnlY a Simple SiIll.ilar to N. 8Ordida, bUt at least one species, No structure is present basally in Achaetothorax, al- hispanica,isnotso. Female sternite 4 is undividedin Vol. 6, No. 3-4, September - December, 1992 157 thisgroup(i.e. LheplesioulOrphicstal:eofcharacl:er21 which suggests that N. Ii igipennis is more closely is retained). relatedtoN.lacteipennisthantoN. jiLlvipennis. The The oostalis group IS a large Holarctic clade WIth WIng IS also white m N. scripta, mclicating that this onlytwo speciesinthe Nearetic Region, the Aretic No character is subject to homoplasy within the genus yukOnens18 and the Holaretic N. {Umlpenms. These ThescnptagroupmcIuClesN. mexunna,n. sp.,N. species share two apomorphies with a number of scripta (Malloeh), andN. singusta, n. sp. This group otherPalearcticspecieswe haveexamined,including is restricted to the New World, and is primarily N. wstalis. The male cereus is raised posl:eriorly, wesl:em: NearcLic, with one species known from the forming a knob or ridge (character 16), and the Neotropical part ofMAXiM, Synapomorpbies for the hypoproct has a characterIStic shape and a separate scnptagroupmcludetwocharacterstfiatappeartobe bifid selerite associated with it anteriorly (eharaeter unique to it: the shape of female stemites 6 and 7, 17). Thetriangular,anteriorlyacute,bareareaonthe which are bare and shiny anteriorly, with a deep Wiamay be a synapomorphy fOI a group within the transverse groove (character13); and the presenceof oostalis group that includes N. fumipennis and N. a comblike row of 5-8 stout, straight bristles on the yukonensis. The broader, anteriorly more truncate posteroventral part of the surstylus (character 14). shape in the scripta and frigipennis groups and in Another probable synapomorphy is the large size of Ru:JUil'dina isprobablypleSlomorphic. Female stem­ the bare area on the anepisternum (clUiriiCter 15), ite4isquadripartite(eharaeter21.3)inN.{umipennis, although like the hens mierotriehial pattern, this N. yukonensis, and a number ofother species of the character is also highly subject to homoplasy in the oostalis group, which may be a synapomorphy for Copl'Omyzinae. Within the group, tv... mexioana and these species within the group N scripta are probably the more closely related 'Ilie{rigipenn18 groupISpnmarilyeasternNearc­ spec1es (Fig. 3F). This hypothesis is supported by tie. It includes £'\7. frigipennis, N. (ulvipennis, N. eharaeter21,state 1, thesplitoffemalestemite 4intD ku:teipennis, and N. triglabro, which share the de­ two sclerites. The breakup ofsternitesis commonin rived states of characters 18-20. The apomorpbic females, butusuallyoccursindifferentwaysindiffer- state ofcharacter18islessobviousinN. lacteipennis entspecies. We hypothesizethatthetripartitecondi- than m the other specIes, andthatofcharacter 19 IS Bon (state 2) m]V. fUlmpenms andthe quadripartite less obvious in No triglabra, but is especially pro- condition (state 3) in many species of the oostalis nounced in N. j'rigipennis and N. fiilvipennis. Be­ group areindependentlyderived. Insomespecimens cause the female of N. trigZabm is unknown, the of No yukonensis, n. sp., the lateralpairofsternitesis statusofcharacters 22-25is uncertain. Eachmaybe reduced, resulting in a secondarily bipartite condi­ a synapomorphyforthe frigipennis group, oronlyfor tion. N frigipennis, N full1ipennis, and N lacteipennis as Norrbomia scripta, like at least some species of is showninFig. 3D. Similarly,the significance ofthe the fT'igipennis group, is kIeptoparasitic (character kleptoparasitiehabit(eharaeter26)inthelatterthree 26). Like N. lacteipennis and ~v. frigipennis, it also species is also uncertain, because the biology of N. has white wings (character 30)that often are associ­ trigtabm is unknown. Kleptoparasitism also occurs ated with this habit. Ceroptera Macquart in N. scripta in the scripta group. (Limosininae),theonlyothergenusofSphaeroceridae ThemonoplijilyOltllecladeincludingN frigipennis, that is kleptoparaSitiC on scarabs, also has white N. fulvipennis, and N. lacteipennis (Fig. 3E) is sup- wings. Thisraisestheinterestingquestionof'\\rhether portedbytwo male characters, the extreme lengthof kleptoparasitism originated independently in differ­ thebristlesontheposteroventra1partofthesw'Stylus entlineagesofNorrbomia,ornitoriginatedonce and (character27)andthebilobedornotchedshapeofthe was lost in N. singusta. Although it seems unlikely cercus (character 28) Of these three species, N that this complex behaVior should 6riginafi\ twice in frigipennis and N. fulvipennis are probably more the same genus, furLherinformation on the behavior Closelyrelatea: TheyaretheorilyspeciesofNorrbOmia ofthe poorly known species, plus betterresolution of withthe pleuraof female abdominal segments 6 and the I'elationships between the frigipennis, scripta, 7 striate (character 29). They also have a more and oostalis groups are requiredto pursue this ques­ similarly shaped male cereus and have the setae on tion the base of the cereus and bordering area of the epananumSliglitlymore denselyclusteI"edthaninN. 1acteipennis (see character 19). This hypothesis is contradicted by the color ofthe wing (character 30), 158 Insecta Mundi

Key to the Nearctic species of Norrbomia Frons with eiungate bare area wIte! iOl to ante!iOl ocellus. Male hind tibia with or without row of 1. Tarsi, other than tarsomeres 1 and 2 of hindleg, long thin posterodorsal setae. Feznale stemite 4 white; ~mainderoflegs brown. Wing membrane undivided or divided. Wing membrane brownish and all veins white. Female sternite 4 smgle, ...... 6 about half as wide as sternite 3 .frigipenms (Spuler), Flonda to Massachusetts. 6. Hind femur not strongly swollen. Male hind tibia 'l'ajOjij binW1i arpale brown Wing variable E'AiIJale withonlyshortposteredorsal setae. Femalester sternite4 dividedintopiecesoraswideassclerite nite 4 divided into 2 widely separated parts (Fig. 3 2 80)...... mexioona, n sp., Cbiapas, Mexioo

2. AnepislenIulIl with nonpruinose shiny area small (Fig. 21), its dorsal margin closer to level of s rni Unl~e g:~ .. anteroventral OOlnel of anepiste!nwn than to singustn., n &p., Western North America level ofventral ma~of s~iracle. Fronsentirel,Y ptuinose, or (11," fitiVipe1uzi.s) withsmaHbw'ewea 7. Genalpruinositydense,wbitish, stronglycontrasted lateral to_~teriorocellus not much larger than with bare area, which is narrow (its height at ocellus. Wing membrane transparent or with narrowestpointapproximatelyhalfthatofpnilil- brownish tint 7 use area) but slightly expanded at anterior mar­ Anepistemum with nonpruinose shiny area large gin (Fig. 19). Only 1 (prescutellar) pair of large, (Fig. 24), its dorsal margin dorsal to level of well differentiated dorsocentral bristles . ventral margin of spiracle or at least closer to it ...... 80ttlida (Zettersledt) than to level of anter(wentral corner of Widespread Holarctic, south to Guatemala. aneplstemum. Frons Wlth bare area Iateral to Genalprumositylessdense,notstronglyoontrasted ocelli at least aslargeasareabetweenocelli(often with bare area, which is broad and truncate or of difficult to see lU N. 1aCteipenm,s (Fig. -I». WIng acutely tapered anteriorly. 8 (1+2) pairs large, membrane often opaque whitish 3 well differentiated dorsocentral bristles. 8

3. Anepistemum with dorsal margin of bare area 8. Genal barearea tapered, acute anteriorly(Fig. 16). ventral to level of dorsal margin of spiracle (Fig. Frons entirely pruinose. 3-5 interfrOntal setae. 28) ...... 4 Female stemite 4 divided into 2 or 4 parts. Anepistemumwithdorsalmarginofbareareaator dorsal to level of dorsal margin of spiracle (Fig. 24) 5

4. ocellus. 6 or more mterfrOntal setae. F'emaIe roxunately 2.0 t-es ge ~Pala£~al 8ternite4diVided into3parts(Fig 62) Hypopmct narrower than height of pruinose area on gena, slightlyconcave anteriorly, without separate,bi­ approximately VIO of height of antennal furcate process (Fig. 60)...... titlvipennis, n. sp. flagellomere...... triglabra, n. sp., Kentucky. Eastem and central United States to eastern \\Gng membrane opaque whitish. Eye small, its Mexico. height approximately 1.5 times gena height. Parafaeial broader than height of pruin.ose ama 9. Posteromedial part ofmale steruile 5 pale, densely on gena, at least VB of height of aniermal microsetulose. Posterior margin of paramere flagellomere (Fig. 10) ..... lacteipennis (Malloch) evenly convex (Fig. 54). Male cercus in laleral SOllth oontral United States to northeastern view (Fig. 53)with dorsal part slightly produced. Mexico. Armofbifurcaleprocessofhypoproctsubequalin length to middle part of hypoproct (Fig. 72) ...... 5. F'I'Ons prumoseantenortoantenoroceIlusorWlth ...... yukonensis, n. sp., Y1:1kon. bare area not much larger than ocellus (Fig 8) Posteromedial part ofmale sternite 5 not differen- Malehindtibiawithrowoflongthinposterodorsal tiated from rest of sternite. Posterior margin of setae. Female sterilit.e 4 diVided (Fig 85) Wing paramere deeply notched, thus appearing sinu membrane usuallywhitish..... scripta (Malloch) ate(Fig. 33). Malecercus in lateralview(Fig. 35) western North America, Mexico. :::=~:~sJ;~::~.:e~~:: middle part of hypoproct (Fig 71)...... fUmlpenms (Sterihammar) Northern Holaretie, Rooky Mountains. Vol. 6, No. 3-4, September - December, 1992 159

NOllbomia frigipermis (Spoler) (Figs. 26, 28)densely setuloseposterolaterally,poste­ new combination rior setae longer than surstyl.USj anterior part of (Figs. 4,14,20,26-28,56-60) surstylus short, WIth cannate, antenorly pomted ventral lobe Cereus with dense patch of setae poste1'Odorsany;posteroventr8IIyshortsetose,biIobed, Borboros ~ Knab 1915:40. outer lobe narrow; inner lobes of eelei 8epm:ated by

contiguou . Param.e (Fig. 27) bro near middle, (USNM). COpromyza (Borborillus) frigipennis: Richards ~:~:~;;~ari::~7a:;e:~=p=: 1965:720; Steyskal.1971:477. undivided, apieallywith single, medial, dorsally pro- 1307 bo7'illus fl"igipennis. Sivinski 1983.419, 1984.232. jected lobe; distal part of distiphallus larger, with. rounded disl:a1 ventral lobe and paired, dorsoven- Description. Body length 1.6-4.5 mm. Body color dark brown to black.; antenna and middle of race reddish; coxae, femora, tibiae, andfirst 2 tarsomeres

dinally striate (Fig. 58). Talps 6 and 7 (Fig. 56) In rvlbnssaldistance; 8I"l Immnn. asal complete anteriorly and trilobed posteriorly, middle width olarists. Frons(Fig. 4)pruinose, with medium part very pale. Terglte 8 biparttte; epiproct (Fig. 60) sizedbarearealateraltoocelli,onlypartonnonrugose eutieIe extended postei'Olaterally from posteriorooeI obviousl~shin~. Ius Interfrontal bristlesin6-8 small elongate,bipartiteexeeptalonganteriormargin,with single transverse row ofbristles. Sternite8 bipartite, 1.~ gen~h bl~ with dark put!teromediallobes. and tbirflies. nonmicrotrichose area broad, tnmcate anteriorly or This specIes IS kleptoparaslfic on the dung caches of Its anteriormargIn concave. T1wrax: KatepiSternum dung burying searabs, and has also been oo1leGted bare below level of dorsal setae; anepisternum (Fig. clingingtothe adultbeetles(Knab 1915, asBorbo1U8 21) withtinyisolatednonmicrotrichose areaimmedi­ species -witll milk-white wings"). Sivinski (1983, atelyan:teroventraltospiracle,mainnonmicrotrichose 1984) describedits biology. He found that adult flies arearelativelysmall,itsdorsalmarginwellventralto interceptadultscarabsandusuallyclingtothe dorsal level ofventralm:argi.n ofspiracle; pleuronotherwise surfaCAoftbebeetleon theposUlriorpartofthethorax lightly proinose; scutum and scuteJlum uniformly, orthe antenorpart ofthe eIytra: BOth sexes accom­ heaVilyprumose. Wmgmembranetransparent,WIth panythe beetles underground, andoviposition takes light brown tint; all 'lems pale brown; seoond costal place in the dung cache. Specimens have been ob­ sector2.0-2.2Xaslongasthird.Forelegsetose,brown; servedinassociatiOIlwithPhm!eUSvi7u:kx(MacLeay), tarsi pale. Midleg brown, tai-si pale; first tarsomere P. igneus (MacLeay), Boreocanthon depressipennis withslightlyenlargedventral bristles. Hindleglong- (LeConte), Onthophagus spp., Canthon pilularius setose,brown; tibiawithsmall anteroapicalspurand (L.), and Co imitator floridanus Brown [pl"e'oously large apioo'lentnU spur; femur with 3 erect dol'sal misidentifiedas C. viridisinKnab(1915)accordingto bristles on distal h.8lf. Postel'OdorsaI setae of dista1 Steyskal (1971»). half of male hind tibia longer than tibial width. Sivinski (1983) found thatthe sex ratio ofboth lab Pulvilli D.81TOW, pads widely separated. Male abdo­ rearedandfteldcollectedspecimenswas significantly men:Syntergite1+2uniformlypruinosebl\JWll.Mem- female biased. The extreme size variation in N braneofsegments3-5with setaeon scIerotized bases. {rig7,penn/,S does not reflect sexual dimorphism St.enute 5 mmple, darkly pIgmented along posterior (Si'Jinski 1983). Si'Jinski (1984) diseusses the rela margin. Epandriumwith dense, shortsetaedorsally; tionship betweenmale size andreproductive success laterally with 4-5 long bristles, dorsolateral bristle in Lhis species. longest. Surshlus (Figs. 29, 31) seLulose Norrbomia frigipennis, N. lacteipennis, and N. posterolaterally,posteriorsetaelongerthansurstylus; {Ulvipennis form. a monophyleticgroupcharaetenzed antenar part of surstylus elongate, WIth smatl, can­ by similar male genitalia, pale tarsi, and several nate anteriorly rounded ventral lobe. Cercus with attributesofthefemaleabdomen,includingthefourth densepatchofsetaeposterodorsally;posteroventrally sternite broken into a median remnant and several shortsetose.bilobed,outerlobebroadandshort;inner lateral pieces. lobes of cerci separated by deep apical notch, cerci posteromemallycontiguousoverdorsalhalf. Paramere Norrbomia fulvipennis, new species (Fig 30)broadnearmiddle,tapered roan acute apical (Figs. 5,11,15,21,29-31,61-65) process. BasIphaIlus WIth long, narrow epIphaIlus. Basal selerite ofdistiphallusundMded, apieallywith single, medial, dorsally projected lobe; distal part of disLiphallus larger, with rounded distal venLrallobe or rus Stngu7"tS:iVIDS 1983:419(mIS-Identifi­ and paired, dorsoventrally flattened, distal dorsal cation) lObes Female abdomen' Stemite 4 (Fig 62) diVided into 3 separate sclerites, selerites weak orbroken up Description. Body length 1.6-3.5 mm. Body color on some specimens; sternite 5 entire but pale medi­ dark brown to black; antenna and middle of face ally. Hearon of segment 6 I ugose anteriorly and reddish; coxae, femora, tibiae, and first tarsomere of longitudinally striate posteriorly, pleuron 710ngitu­ :::go~wi:rr~~t:gr: dinaJly Rtriafu (Fig. 63). Tergites 6 and 7 (Fig. 61) completeanteriorlyandtripartiteposteriorly,middle greaterthan intervibrissal distanoo; aristal hairs roo part very p8.Ie. Tergite 8 bipartite, epiproct (Fig. 65) ax basalwidthofarlsta. Frons(Fig. 5)pruinose,with elongate, emarginate anteriorly, produced posteri smaH, indistincL, nomnicrolrichose shiny spoL beside arly, bare except for 2 bristles. Sternites 6 and 7 elongate,bipart1teexceptalonganteriormarein,with single row of bristles near middle. Sternite 8 tripar­ luteou8onmedialhalf,blaekonouterhalf.Eyeheight tite, Willi aark posteromedial lObes; median part roo 2.5Xgenalheight. Gena(Figs. 11,15)darkreddish narrow and weakly sclerotized. Hypoproct emargin- brown, microLrichose anteriorly and fairly broadly ate and bare anteriorly, setulose posteriorly. Sper- Vol. 6, No. 3-4, September - December, 1992 161

Flguree4-9. DorsalviewofheadofNorrbomiaspp. 4,N. frigipenni8; 5,N. fulvipenni8; 6,N. fumipenni8; 7, N.lacteipenni8; 8, N. .cripta; 9, N. 8Ordida. 162 Insecta Mundi malheca (Fig. 64) subspherical, flattened at lop and ·VERACRUZ. Allende, vii.1944, M. Guerra (1m bottom, with large basal apodeme; neck 0.75-1.20 AMNH). times as long as mmn part. Distribution. Nebraska, Miehigan and Massachu Holotype. male, VNITED STATES. FLORIDA. setts to Florida and Veracruz, Mexico. Collier Co., Seminole State Park, 28.xii.1979, 8,A, Marshall (CNe). Comments. Norrbomia fulvipennis is widely dis- tributed in eastern North America, but unlike the Paratypes.VNITEDSTATES. ALABAMA. Kushla, closely related N frigipenni.s, does not seem to be "Oc.'24", Sturtevant CollectIon, (1m USNM). abundant anywhere. Most speCllIlens have been FLORIDA. AlachuaCo., Gainesville,2O.i....1967,light taken on dung or in dung baited traps. Sivinski trap, W.W. Wirth (1m VSNM); near Gainesville, on (1983)record.edthisspecies(asBorborillussingularis backofOanthtmpilumris,8.v.1981,J.Sivinski;Iligh- Spuled as phoretic wI adults of the dung-rolling lands Co., Highlands Hammock State Park, orange scarab Canthonpilurarius (L.). Sivinski also reared grove, pig dung, 14-18.Vl.1982, Woodruff & Rench mdiVlduals from a dung caChe of the nocturnal, bur­ (5mlfFSCA); TekePlacid,2-3 xi.1983,pigdung,R.E. rowing scarab Copris minutis (Drury). The larger WOOdrUff (5m5fFSCA); Al'ChbOld BiologiC8I Station, "Borbo1U8" species "with smoky wings" reported by "'8GUumedfromdung,17.iv.1989,S.A.Marshal1(lm); Knab(1915)asridingonCcmthonimitatorfloridamts Marion Co., Ocala National Forest, 12-18.vi.1983, (as C. viridis, see comments under N. {rigipennis) 7.vi.1984, and 15-16.iii.1984, RoE. Woodruff (lm3f); might be N. fuluipennis. The flies from Missouri OrangeCo., E Orlando, 2-3.vii.1982, sand pine scnJb, reported by Moulton (1880) are probably N. scripta, RE. WoodrUff (1m); Okaloosa Co., BlackWaterRiver not this speCIes as suggested by SivinSki (1983, asB. StateForest,FloridaA&Mreseareh.station,pigdung, singularis). 15-17.vi.1978, L. Stange(1m); PolkCo., 7miN 14, 2­ This species is closely relatedto N. {rigipennis and 3.xi.1983, pig dung,RE.Woodruff(lm);SarasotaCo., N. lacteipenni6; from whichitcanbe separatedbyits Myakka River State Park, 21.v.1982, pig dung, RoE. brownish wings. WoodrUff (lmlf). GEORGIA. Towns-Umon Co., Mt. 4:2~ 23.ix, 2tx.1952,trap6, H.R.~ Norrbomia fumipennis (Stenhammar), ='W V,V NM); Ra un Co., RabunBald, 24.lX, new oombination 25.x.1952, trap 6, H.& Dodge (8mBfW8U, USNM), Chatham Co., Savannah, 14.v.1954, privy trap 84, (Figs. 6, 16, 22, 32-34, 66-71) J.\V.Kilpalrick,(2m4fWSU,USNM)jFolkston, 14mi N, dungtrap, 1O-18.iv,l 989,.1 Swann(5m2f) MARY- Copromyza fUm~pennZ8 Sterihammar 1855:352. Lee- LAND. MontgOmery Co., v.1959, light trap, A.A. totype male [here designated] (NBS). S\VEDEN. Hubert(1mUSNM).MAsaA&GHUSErrS.DukesGo., Lapponia Lulensi (=Lule, Lappmark), C.H. Naushon I., 2.ix.1923, A.H. Sturtevant (1mVSNM); Nonamesset I., 24.vi.1923, ahem> dung, 7.vii.1928, A.H. SturtevantCollection(4mVSNM). MICHIGAN. I.htingston Co, George Reserve, 17 iX 1949, KK. Bohnsack, (1m USNM). MISSISSIPPI. Hom I., Fort Chimo [with following labels: "Ft. Chimo, 1O.v.1944, light trap, E.A. Richmond (If VSNM). La:b:r, 7.22", "LM. Turner Collector"j red "Type NEBRASKA•. ThomasCo.,NebraskaNationalForest, No. 14950 V.S.N.M."; and "Borhorus articus 2.5 mi W Halsey, 16.vii.1967, H.B. Leech (If CAS). Malloch"]. new synonym. NEW YORK. Riverhea

15e. ------S'U '0 16" S 99999 P 99919

Figuree 1()'13. Left lateral view of head of Norrbomia 8pp. 10, N. Iacteipenni4; 11, N. fulvipenni4; 12, N. scripta; 13, N. sordida. inneroroitalaetulae.Eyeheightoo.2.1Xgenalheight. bristles on distal half. Poaterodorsal brietles ofmale Gena(Fig. 16)dark,broadlypruinoaeanterovent.-ally, hind tibia slightly shorter than tibial width. Pulvilli nonpnrinose area tapered, acute anteriorly. Thorax: narrow, pads widely aeparated. Male abdomen: Katepiatemum bare below level of dorsal aetae; Syntsrgite 1+2 uniformlypruinoae black. Membrane anepisternum (Fig. 22) with tiny, isolated of aegments 3-5 bare or with 1·2 minute aetulae. nonmicrotrichoae areaimmediately anteroventrslof Sternite 5 large, dark; posterior margin prominent spirscle,mainnonmicrotrichoaeareasmall,itsdorsal and weakly concave posteromedially; ventrsl part of margin well ventrsl to level of ventrsl margin of aternite 6 thickened, dark, with conspicuous concav· spiracle; pleuron, including meron, otherwise en­ itybehindpoateromedialpartofaternite5.Epandrium tirely lightly pruinoae; scutum and scutellum uni­ withaparae, shortsetae dorsally; laterally and poste­ formly, heavily pruinoae. Wmgmembrane transpar­ riorly with a few longer bristles. Surstylus (Fig. 32, ent, with light brown tint; all veins brown; second 34) aetuloae posterolaterally and vent.-ally, longest oosta1 sector 2.8-3.0X as long as third. Forelegblack, bristles much shorter than surstylus length; with knees sometimes reddish. Mid tibia dark, fn-st compact group of bristles posteromedially, anterior tarsomere with slightly enlarged ventrsl bristles. part of surstylus broad, elongate, mostly bare, with Hindleg short-aetoae, dark; tibia with single, very small, carinate, anteriorlyroundedinnerventrsllobe long apicoventrsl spur; femur with 2 erect dorsal andaecondsmallinnerlobenexttoapexoflargeinner 164 Insecta Mundi lobe. Cercuswithrounded,bareposteriordepression, FOUNDLAND. Bay St. George, 10.vii.1915 (1m apical partofcereusthus scoop-like Cew divided by AMNII) NORTHWEST TERRITORIES Salmita broad,deep,trianguIarcIeft.Paramere(F'ig.33)broad Mines, 64"05'N, ll'15'W, Vl-Vlll.1953, J.G. ChiIlCOtt near middle, posterior surface strongly smuate be (1Om2fGNG);Tuktoyaktuk,8 12.vii.1971,D.M.Wood yond middle, tapered to digitate apical process; 5 (3f CNC); Chesterfield, 9-24.viii.1950, bred from. pu­ setulae along posterior surface and 2 preapieal setae paeinLemmusbun:ow,J.RVoekeroth(10m7fONC), anteriorly. Basiphalluswithlong,n8lTOwepiphallus. Aklavik., 2.vii, 5.viii.1931, Bryant(lmlfCAS); Masik Basal scleriteofdistiphallus undivided, apicallywith It, Banks I., 17.vii.1968, G. Shewell (1m CNC); Es­ single, medial, dorsaJ)y projected lobe; distal part of kimo Point, 16 vii 1950, G G. Dilabio (1m); Oyde, distiphanus larger, Wlth rounded distal ventral lobe Baffin Is., 2.Vlll.1958, G.E. Shewell (2fCNC); Spence and paired, dorso'lentnilly flattened, distal dorsal Bay, 13.vii.1951, J.G. Ghilleott (1mlfGNC); Muskox lobes. Female a1:x1nmen: Stemite 4 (Fig. 67) divided L.,64·45'N, lOS'lOW, 11-25.vii.1953, J.G. Chillcott into 4 separale scleJ:iles; slenrlle 5 entire but with (7m5fCNC);FordLak.e,6S·11'N.1OS·10"N;27.vi.1966 sma]] anteromedial notch. Pleura ofsegments 4 and (If CNG). QUEBEC. Ft Chima, 29 vii, ".M Turner 5(Fig.68)Wltha few largesetulae,someonsclerotiZed (lfpara1eCtOtype ofB. artwus USNM); Indian House bases. Tergites 6 and 7 (Fig. 65) oomplete, dark, T eke, 8.vi-5.viii.1954, W.R. Riehards (8m7f CNG); tripartite posteriorly. Tergite 8 complete posteriorly, Payne Bay, 18-00.vii.1954, R. McCondochie, W.R.M. dark, shiny. Epipreet (Fig. 70) long·setulose, with 4 Mason, B.B. McDougall (3m4f ONO); Sugluk, larger bristles. Stemites 6 and 7 bipartite except 2.viii.1954, H. Huckett (7m2f CNC); Pt. Hanison, alonganter.iormargin,withsinglerowofbristlesnear 9,lO,25.vi.1949, D.P. Williams, P:J. Lachaine (lmlt middle Stemite8 bipartite, dark, posteromedialcor- CNC); Ot Whale River, 7,8,15 viii 1949, bred from ners shiny and pr<>Jected ventrally; some mdiStinct pupae in nest of M~crotus, J.& Vockeroth (2Di2f sclerotizationin gI'OO'Ja betv.'OOn hal-Jas ofsternite 8. CNG). YUKON. Riehardson MtDB., flood plain of Hypoproctemarginateanteriorly,withbifurcatescle­ small creek, 5.vii.1987, S.A. Marshall (1m); 67'58'N rile in emargination. surface of hypoproct setulose, lOO'29W, 1-5.vii.1987, dung traps in wet tundra, S. bifurcate sclerite not setulose. Two small vaginal Marshall (11); Otter Lake, 100'25'N 62'OOW, 4000', sclentes VlSlble on cleared specnnens. Spermatheca 16022.vii.I960,J.E.H.M8rtiIi,E.W.ROCkburne(IIm7f (Fig. 69) subspherieal, with large basal apodeme CNG); Hersehel Is., 1020.vii.1963, C.D. Bird, J.B. extending almost length of main part; neck 3-4X Waterhouse (3m2f CNC); Tombstone Mtn. camp­ longer than main part. ground on Dempster I1wy, musluoom traps, 5- 11.vii.1985,SAMarshall(1m); NorthForkCrossing, Material examined: Leetotypes of C liLmipennis mj43pAAlPlt Rd ,3!500',3Vii 1962,R:E Leech(lmlf and B arnclls (see synonymy) SWEDEN "Lp in ", CNC); Firth. lliver, 24.vi,17.viii.1956, E.F. Cashman "Bhlri" (lDl2fparaIectotypes ofC. fUmipennis NRS); (2CNC);HainesJet.,2.viii.1958,Mason&Hughes(If Norrbotten, 16 km 'II Messaure, 16.vi.1967, hV. ONO). UNITffi) STATES. ALASKA. Umiat, 3- Knutson(IfUSNM);'Ijuonaj,28.vii.1926, 3O,vii.1918 24.vii.1959, R. Madge (9m8f CNC); Ulnalakeet, 4­ (3m ZIL; USNM);Jebren,j, 240vii.l918 (lmlfZIL); II- 22.Vij,1961, R Madge, B SHAmming (2mlf CNC); a ker, 28.vi.1922 (1m ZIL); Nuolja, 25.vii.1926 (1m CapeThompson,25-29.vii.1961,B.S. Hemming(3m3f ZlL); Villis:ta, bJii.1939(1mZIL); Abisko, 19.vii.19IB CNC); N8kIlCk, 6,18.vii.1952,'N.R. Mason(2mGNG); (IfZIL). CANADA. BRITISH GOLUMBIA. Tunjony St. Paul Is., 14 22.vii.1980, Adler(1m USNM), same, L., 58'26'N, 132°45W, 2O.vii.1960, flowers, grasses 10.viii.1920, G.D. Hanna(2m!CAS, USNM); St. Paul and sedge, 3200', R. P.ilfrey (1m CNC); Lisadele h, Is.,fwmTlrlck-billedMurre, 22.vii.1980(1m); Sealer's 58'41'N, 133'04W, 11.vii.1960, 4000', R. Pilfrey (If Is., 10.vii.1965(lm31); IsabelPass,mi206Richardson CNC). MANITOBA. Churchill, 15 Vi 1930,0 Bryant, Hwy, 4500', 18 Vii 1962, RoE Leech (1m CNC); 17.vii,1930 (2m CAS), 30.vi.1952, 26.vi,1948, RichardsonHwy., mi 206, mammalrunsnearbeaver 3.viii.1952, 25.viii.1950, 26.vi.1948, 18.vii.1952, pond, 19.vi.1987,S.A. Marshall(lm1f); MillerHouse, vm..1962, J.G. Chilloott, G.B. Shewell (3m6f eNC). Steese Hw:v, 10.vii.1937, C.B. Philip (if USNM); LABRADOR. Ungava Bay, 24.vii, L.M. Turner (If EagleSummit,54miSWCircle, 1105m,caniontrap, paralectotype of B articus IISNM); Hebron, 60 13 Viii 1984, S &J Peck(lf) COIDRADO Love1alld 1O.vii.1954, J.F. McAlpine (7m5f CNC); Nutak, Pass, 12,000', 7.viii.1961, J.G. Chillcott (lm4fCNC); 23.vii.1954, J.F. MeALpine (2Dl3feNG); G8ltwright, Boulder Go., Gorona Pass, 10,600', 1l.viii.1961, 19.vii.1955, E.E. Sterns (1m CNC); 10mi up Saglek 1l.viii.1955, J.G. Chillcott (lm2f CNC); Clear Creek Fjord, 3.viii.1954, J.F. McAlpine (6m CNC). NEW- Co., Mt. Evans, 12,800', 24.vii.1961, B.H. Poole (If Vol. 6, No. 3-4, September· December, 1992 165

Figureo14-19. Left senaofNorrbomia spp. 14, N. frigipennis; 15, N (uluipennis; 16, N. (umipennis; 17, N. lacreipennis; 18, N. scripta; 19, N. BOroida. 166 Insecta Mundi

CNC), Gunnison Co., CotLonwood Pass, 11,000', Norrbomia lacteipennis (Malloch) 29.vii.1961, J.G. Chillcott (If CNC); Lake Co., Inde- nev.' combination pendence Pass, 12,100', tundra, 31.viI, 7.Vlll.1961, J G Chillcott (2m4f CNC); Larimer Co , Cameron (Figs. 1-2, 7, 10, 17, 23, 35-37, 73-78) Pass, 11.viii.1964, G.F. Knowlton (If CNC). NEW MEXICO. Taos Co., 1.7 mi BE Tres Ritos, 8500', Borborus lacteipennis Malloch 1913:366. Lectotype dung 3-5.vii.1972 A.Newton~lf). UTAH. Duchesne male [here designated] (USNM). MEXICO. ~ Co/Mirror Lk., 10800', 8O.~~12.viii.l979, S.&J. TAMAIIIJPAS Tampico, "290 1 EA Schwarz" (withfollowinglabels,"TampicoMex29.12","E.A. Sehwarz Colleetor", red "Type No. 14949 13.vili.1957, G.F. Knowlton (1mlf WBU); Carbon U.S.N.M.", "BorboruslacteipennisMalloch", red- Co., Lake Marie, 20 km. W Centennial, 3230 m, bordered "Holotype Borborus lacteipemris Mall. sweep, 1.viii.1973. P. A1Il8Ud (3m7f CAS, USNM); ::YS~l~O" . Sublette Ct;Wind River Mts., 20 mi NE Pinedale, Bor rus or rillus) lacteipennis: Spuler 1925:9. nr. Nelson ke, 10400-11000',carrion,tundra, 15- BorbOnts (BOrbOrillu s) peltastes Spuler 1925'10 Hoo 23 viii 1979, SPeck (2m10£) 10typemale(USNM).UNITEDSTATES.TEXAS. Mobeste(sic), A.lN. Barber(withfollowinglabels, Distribution. Throughoutthe northern Holarctie "Mobeetie. Oct 9, Tex", orange "PARATYPE Region, occurring south to New Mexico in the Copromyza peltastes Spuler", "AL. Melander mountains of the western United States. Collection 1961", "Copmmyza peltastes Spuler" [Melander's writing], and"Type male, this is the Comments.NorrbOmLafUmtpenms, N. yukOnensLs, realholotype, det. Sabrosky"); Steyskal1971:478 and N. sordida exhibit a number of similarities, (synonymy). such as the setulose epiproct andentirelypruinose Copromyza (Borborillus) lacteipennis: Richards ~ey:~~~q~l'~ frons, shared with other species of Norrbomia not Coprom~(1967'2' found in the New World. Norrbomia fumipennis orhs Pe;s:Riclial'dsl965:720. and N. yukonensis are very closely related to N. costalis (Zetterstedt), with which they share a Description. Body length 1.6-3.5 rom. Body color quadripartite female sternite 4, posteriorly con­ hIowntoblack, heavilypItW:lOSe anddarkerdorsally, antenna~e:J:iand cave male cereus, a charaeteristic SUIstylus shape, lowerfrons and:nareddish; face and the possession of a separate sclerite in an anterior emargination of the hypoproct. These species differ most obviously in cercus shape, the latter species having an elevated posteromediaI corner. The Palearctic species ofthe costalis gtOup are in need ofrevision. Kim (1972) did not report type specimens of C. fumipennis in the NRS, and P.I. Persson (pers. - mc mner 0 1 setti ae. ace ep y con- comm )fnnndnospecimens'mderthisname HOW- cave, uniformly luteous. Eye height 00. 1.5X genal ever, among the series of Sphaerocera species, he height (Fig. 10). Gena (Fig. 17) pale reddish brown, located four specimens fitting Stenhammar's de­ narrowly prtW:lOSe anteriorly and ventIally (extreme scriptionandwithapPI'Opriate labelstobesyntlpes. ventral=n bare~onpruinosearea very broad, The label "Lp.in." is an abbreviation for "Lapponia interior", orlnnerI.appland,whereI dJle, I .appmal'k islocated,and"Bhm"meansBoheman(P.I.Persson, anepistemum Fig. )wi nonpruinoseareamo r­ pers. comm.). atel)' large,itsdorsalmarginslightl)' venttaltoslightly ofventralmar~n Ft. Chima, the type locality of B. articu8, is in dorsaltolevel ofspiracle;pruinosity northernQuebec,ratherthanLabradorasthelabel extended towards or to an roventral corner, par- of the lectotype and the original description indi- tially or ::;'=ly ~sol~ tiny :~nose.area cate. NO.,.,.bomia fumiPennis has been reared from and collected as adults in burrows of rodents scutellum uniformly, heavily pruinose. Wing mem­ (LemmuB and Microtus spp.). brane opaque, whitish; all veins usually whitish, Vol. 6, No. 3-4, September· December, 1992 167

Figures :»-21>. Left anepi8lernum ofNorrbomia 8pp. 20, N. {rigipennio; 21, N. fuluipennio; 22, N. fumipennio; 23, N. lacteipennio; 24, N. 8cripta; 26, N. 80rdida. 168 Insecta Mundi cosLa sometimes and ollier veins rarely pale brown; paralecwLype of B. lacteipennis USNM). UNITED secondcostal sector2.1-2.3X as long asthird. Foreleg 5TATES.WUISIANA.Cameron,7.vij,1905,J.S.Hine lightbrown, tarsipale. Midlegbrown, tarsipale; firSt (Im4f OliSU, USNM); Logansport, 24.1ll.1908, E.S. tarsomere with slightly enlarged ventral bristles. Tueker(2fUSNM). OKLAHOMA. LattimerCo., 500 Hind femur with 3 slightly enlarged erect dorsal W Red Oak, 8-14.v.1977, dung, K W. Stephan (2mlf setae. Distal 4/5 of hind tibia ofmale with 8·10 long, FSCA); Comanche Co., "Nichita Mts. Refuge, 5- thinposterodorsalhairsen. 3X aslongastibialwidth; 7.vi.1979. shortgrass prairie, S.&.1. Peck (lm1D. both sexes willi small anteroapical spur and large TEXAS. Mobeetie, 9.x, A.W. Barber (3f paratypes of apicoventral spur Pulvilli nalTQW, pads widely sepa- B peltaste.s IISNM); Brazos Co, College Station, rated. MaleabdOmen: SyntergIte 1+2prumose, dark 2O.IV.I935, H.J. ReiIih8rd; Burleson Co., reared In brown with blaek strip before posterior mmogin, pos lab, 1978 "from STK" (5m6f USNM); Gah.'Cston Co., teriormargin pale; syntergite 1.7X as long as tergite Galveston, "May", F.H. Snow(3m2f UKaL, USNM); 3. Membrane ofsegments 3-5 wiLh a few slIm seLae Kleberg Co., 24.ix.1970. from Phanaeas diffonnis with dark, sclerotized bases. Sternites 3 and 4 re- Lee., RR Blume (4m14f USNM). duced to a few small patches; stermte 5 diVIded longitudinally. Epandrium with dense, short setae Distribution. From eastern Oklahoma, Texas and. dorsally and a few longer bristles laterally, dorsolat­ Louisiana to norLheastem Mexico. ex al bristle longest. Smstylus (Figs. 35, 37) with dense, long setae ventrally, some setae longer than Comments.Norrbomia lacteipennisisrecognizedby surstylus. Anterior part of surstylus elongate, with its small eyes, sclerite-based pleural setae, and deep small, carinate inner ventral lobe Cercus hilobed, cleft separating the male cerci The pnJinosit¥ pat- mnerlobe elongate, narrow. cercIseparated by deep terns are difficult to see. Like N. fUlvtpenms, N. notch. Paramere(Fig. 36)broadnearmiddle, tapered frigipennis, andN. scripta, N. laeteipeRnisisphoretie to apex; posteriorsurface with scalloped, setose edge on coprophagous scarab beetles. Adults have been llearmiddle.BasilJimlluswiLhlong, narrowepiphallus. collected from Plumaeu.s difformis LeConte. One of BasalparL ofdistiphallus well sclerotized; distalparL the three female paratypes of B. peltastes in the smaller, WIth rounded distal ventral lobes. Female USNM was mIslabelled as the "TYPE", probably by abdomen: Sternite 4 (Fig. 74) completely absent; Spoler or Melander. Spuler clearly stated in the sternite 5 verypale, membranous medially. Pleuraof description that the holotype was a male. Steyskal segments 3-5 with numerous setae (Fig. 75), each (1971) said thaL lIe "labeled" Malloch's male synl;ype arising from separate large, dark, sclerotized base. ofB. lacteipennis "as holotype", butthis designation Terg1tes 1-5 (Fig. 73) complete; tergites 6 and 7 with canbemadeonlyintbeoriginal description. Steyskal's posteriorly concave anterior part and weakly pig- statement is not a lectotype designationbyinference mented tripartite posterior part; tergite 8 divided ofholotype (see Intemation8.1 COde ofZoologicaI No­ medially; epiproet (Fig. 78) with bilobed anterior menclature,Art. 74b), becauseMallochclearlystated process and simple posteriorprocess; bare except for that he had more than one specimen. We hereby 4bristles.Sternites6and?eachdividedintoanterior, designate the male as lectotype to eliminate any bare, shiny, posteriorly concave parL and posterior, ambiguity. paler, posteriOrly tripartite part Sternit.e 8 diiiiaea into2 large,dark,lateralseleritesandpalerridge like Norrbomia mexicana, new species sclerites in the ventral groove between the main (Figs. 38-40,79-83) decites ofst.ernite 8. IIypoproct (Fig. 77) pale, con- cave anteriorly, entirely setulose. Spermatheca (Fig. Description. Bod.~ le~h en. 2.0 mm. Body color 76) subspherical, flattened at top and bottnm, with reddish brown to b ac ; antennae and legs reddish large basal apodeme; neck4-5X as long as mainpart brown;balterstembrown,knobwhitish Hea.d:Aristal lengLh equal to intervibrissal distance; aristal hairs Material examined: Lectotype of B. lacte!Pennis, en. 3X basal width of arista. Frons pruinose, with: holotype of B. peltastes (see synonymy). MEXICO. large, triangular nonmicrotrichose shiny area sur- TAMAUUPAS. Tampico, "29.12", E.A. Schwarz (If

FiIfW'e& 26-37. Male }llerreemw,. Left lateralview ofterminalia; aedeagus and associated structures; posterior view of tenninalia. 26-28, N. frigipennis; 29-31, N. {ulvipennis; 32-34, N. {umipennis; 35-37, N. lacteipennis. Vol. 6, No. 3-4, September· December, 1992 169

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rounding ocelli andextended almost l;o anteriormaI"" anterolateral corn:ers with long, thin lateral lobes. gin of interfrontal plate (on some specimens shiny Beuron ofsegment 3 (Fig. 81) with numerous small area diVIded mto 3 parts bynarrow oblique prumose setae ansmgfrommembranenearstermtes;pleuron strips). Interfrontalbristlesin4 6 pairs;6 8 inclinate of segment 5 with a fe\'\T larger bristles arising from inner orbital setulae. Each side of face dark except sclerotized bases. Tergites 6 and 7 (Fig. 79) weakly alongcarinatemiddle partofface. Eyeheighten. 2.0X pigmented posteriorly, tergite 8 with trilobed poste- genal height. Gena dark reddish brown, pruinose riorpartand weakly concave anteriorpart. Epiproct anteriorly and ventrally (extreme ventral margin (Fig. 63) simple, bare exceptfor 2 bristles. Sternites6 hare), nonpruinose area broad, tnmcate to slightly and 7 divided intoseparate scJerites; anterior scJerite tapered anterIorly. Thorax: KatepIsternum dark, bare, WIth deep transverse groove; postenor nonpruinose except posterodorsal corner above dor- partofstemite 7 "'''eakly pigmented, withtransverse sal setae; anepistemum pruinose only narrowly on row ofbristles. Sternite 8 divided into 2 large, dark, dorsal and posteriormargins, nonpruinose areavery lateral sclerites and paler ridge-like scle:tites in ven- large, its dorsal margin dorsal to dorsal margin of tral groove between lateral sclerites. Hypoproctwith spIraCle; pleuron otherwIse lightly prumose except c1istinct anteromecliallobe; bare meclially, setulose fornlll'I'Owbare areaonmeronjscutumandscuteUum laterally. Cerci short, barely twice as long as wide uniformIy, lightly pruinose. Wing membrane trans­ Spermatheca (Fig. 82) subspherical, flattened at top parent, with light mown tint; all veins pale mown; andbottom, with large basal apodeme; necktwice as second costal sectorca. 2.0X as longas third. Coxae, long as main part. trochanters, and tibiae reddish brown; tarsi paler. Hindleg long-setose, brown; femur with 2-3 thin, HoJotype. Male(CNC), MEXICO CHIAPAS 2 mi S erect, dorsal bnstles; tibIa WIth small anteroapIcal Trilrit8ri.a, 21-24.Vlll.I97I,5IOO',Oak-troPICaldecIdu­ bristleandlarge,emvedapieoventralspur. Hindtibia QUS forest, human dung, ~A... Newton. ofmalewithposterodorsalsetaesubequalinlengthto tibialwidth.Pulvillislllall. widely Beparated. t<>geUrer Paratypes. Same data as holotvlJt:l (l3t'9m GUE. much smallerthan tarsomere 5, covering only basal USNM, CNC, FMNH). MEXICO. CHIAPAS. 11 miE half of claw. Male abdomen: SyntergIte 1+2 WIth Trimtaria, 5200', Vlll.1971, tropIcal deCIduous forest, smallpalepatchbetweentergites1and2. Membrane human dung, A. Newton (2m4f). of segments 3-5 with 4-6 small setae per segment, without scle:rotized bases. Sternite 5 sinIple. darkly Distribution. Known only from Chiapas. Mexico. pigmented along posteriormargin. Epandrium with sparse, short setae dorsally, laterally with about 6 Comments. NOIToomia mexioona is recognized by longerbristles, al1 epandrial bristlesrelatively short. its large nonmicrotricbose area on the frons aDd its Surstylus (FIgs 38, 40) WIth smail, short setose pos­ distinctive quadratemalecercI. ItIS closelyrelatedto terolateralarea,....entrolateralmarginwithsomelonger N. seripta. A female from Sonora, 10mi E ofNavaj0 a bristles;posteroventralmarginwith5-6stoutbristles; (in the USNM) may be this species, but differs from anterior part of sW'Stylus qyadrate. with blade-like the female paratypes in having sternite 5 without inner ventral lobe. Cercus with group of 7-8 setae anterolateral lobes. posterodOrsally; posteroventr8Ily ShOrt sefuse, diirer- gent, apex quadrate, with sharp eorners; eerei sepa Norrbomia scripta (Malloch) ratedbybroad,deepapicalnotch,cerciposteromedially new combination contiguous dorsally. Paramere (Fig. 39) broad near (Figs. 9,12,18,24,41-43,84-88) middle, taperedto acute apical process; anteriorsur­ lace With 16bAs above aDd below middle Bai:riphaJll1s with long, narrow epiphallus. Distiphallusrelatively BorbOlUS scriptus M8110ch 1915:64. Holotype maIe simple, withthin dorsaI sclerite andprominentbasaI (INHS). UNITED STATES. IlLINOIS. St. Jo ventralselerite. Femaleabdomen: Stemite4 (Fig. SO) seph, 17.v.1914, J.R. Malloch (genitalia driedout divided longitudinallyinto 2 small, widely separated and: stuck. to bottom ofmicrovial]. Barham.s (BorhariIbLs) .SCripfJlS· Spnler 1925'11 sclerites. Sternite 5 emarginate anteromedial:ly,

Figw-es 38 - 49. Male ATorrbemia. Left lateral view oftenninalia; aedeagus and associated structures; posterier view ofterminaJia. 38-40, N. mexicana; 41-43, N. scripta; 44-46, N. singusta; 47-49, N. sordida. Vol. 6, No. 3-4, September - December, 1992 171

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Cop7'Omyza (B07'b07illus) scripta. Richanls 1965.7'liJ, present. Anterior part of sW'Stylus elongate, with Steyskal1971:477. smaJJ, carinate innerventral lobe. Cercus with a few Borbo1Us (Borborillus) singularis Spuler 1925:12. setae posterodorsatty; posteroventrally short setose, Holotype male (USNM). UNITED STATES. inflexed to anteriorly directed, tapered angle. Cerci WASHINGTON.KettleFans,A.I.. Melander[with separated by shallow, broad apical notch only. followinglabels: "KettleFalls, aMay'12, Wash.", PW8Dlete (Fig. 42) broad new middle, tapet'ed to "AI.. Melander Collection 1961" [added when acute apical process; anterior surface with small incorporated in USNM collection], and triangular lobe near middle and large, quadrate "Copromyza vitripennis Meig" in Melander's preapical lobe Basiphallus with long, narrow wnting. new synonym. epiph8lIus. Bas8.Ipartofdistiph8lIuswellsclerotized; Copromyza (Borborillus) singularis: Richards distalpartlarger, with rounded distal ventrellobes. 1965:720. Female abdomen: Sternite 4 (Fig. 85) divided into 2 separatescler.ites, also often withrenma.n:t:s ofnriddle Description. Size variable, body length ro. 2 mm. sclerite; sternite 5 entire and dark. Pleura of seg- Bodycolorreddishbrowntoblack; antenna,legs, face ments4 and5 (Fig. 86)W1th1-3 distinctsclentes,each andgenaluteous; hal.terwhit.ewithpalebrownstem hearing 1-4 short setulae Tergites 1-6 (Fig. 84) oom- Head: AristaI length equ8.I to intervibrisS81 diSt8ri.Ce; plete, tergite 7 we8.k1y pigmentedposteriorly; tergite aristal hairs 00. 2X basal width of arista. Frons 8weaklypigmentedmediallyjepiproet(Fig.88)simple, pruinose, withmediumsizednonmicrotrichose areas producedposteriorly, bare except for 2 bristles. Ster- lateraltoocelli (Fig. 9), onlypartonnonrugosecuticle nites 6 and 7 each wVldedmto antenor, bare, shiny, posterolateral to posterior ocellus obviously shiny transversely concave part andpaler, setose posterior Interfrontal bnstIes m 5-6 sm8lI Pllll'S; 7-9 mcfiriaf.e part. Sternite 8 divided into 2 large, wk, lateraI inner orbital setulae. Each side of fare luteous on seleritesandpalerridge likeseleritesinventralgroove medial half, black on outer half. Eye height ca. 2.0X between main sclerites of sternite 8. Hypoproct pale genal height. Gena (Fig. 18) dark reddish brown, andbare anteliorly, setulose alung posteriorlIUU'gin. narrowly microtrichose anteriorly andventrally (ex- Cerci very short, less than twice as long as wide. ~e~n~m~nbare1Mnmicromroosearea Spermatheca (Fig. 87) subsphencal, flattened attop very broad, truncate or slightly tapered anteriorly. and bottom; neck longer than main part. ThOrax: Katepisternum b8.l'e ventral and anterior to dorsal setae; anepisternum with nonmietotriehose Material examined. Holotypes ofB. scripta and B. arealarge,itsdorsalmarginapproximatelylevelwith singularis (see synonymy). CANADA. SASKAT­ orslightlydorsaltodorsal margin ofspiracle; pleumn CHEWAN Cypress Hills, 26 v 1955,.1 R Vnckeroth otherwise lightly pruinose except for narrow bare (2mCNC) MEXICO. HIDALGO ElChicoNat. Park, area on meron; scutum and scutellum uriifOriJily, 1 mi S El Chico, 8400', 16.vii.1971, D.H. Newton(1m heavilypruinose.Wingmembraneopaque,whitishor USNM).JALISCO. 7miNTequilla, 17.viii.1960,P.H. occasionally transparent; all veins usually whitish, Arnaud,Jr. (1m CAS). NUEVO LEON. E slope Cerro costa sometimes and othel veDIS rarely pale brown; Potosi, 9200', Pine, human dung; v.1971, A. Newton second costal sector1.8-2.0Xas longasthird. Foreleg (2f). UNITED STATES. ARIZONA. Cochise Co., brown, tam pale Midleg brown, t8iSi pale; firSt ChiricahUaMfs ,6miSEPortaI,41&J',ix.197'~aesert tarsomere with slightly enlarged ventral bristles. grassland, human dung, A.. Newton (4Oml2f GUE, Hindlegbrown;femurwith2 slightlyenlargeddorsal MCZ, CNC); Chiricahua Mts., 7000', 15-21.vii.1978, setae; hind tibia of male with 2-6 long, thin dung, O. Kukal (6m); Coronado NationalMonument, posterodorsal hairs on distal half; hind tibiain both 11-15.viii.1984, dung,B.V. Brown(3m3f). SantaCruz sexes Witb small anteroapical spur and large CO , PatagoDiaT$lie StatePark; 9-11 vlii 1984; dUng, apicoventralspur. Pulvillin8lTOW, widelyseparated. B.V. Brown (15m16f); Santa Rita Mtns., Madera Male abdOmen: Syntergite 1+2 uniformly pruinose Canyon, 5500', ix.1970, human dung, A Newton brown, 1.7X as long as tergite a. Membrane of sag (2m), Made!a Canyon, 14.\'.1977, dung, J. Helava ments 3-5 with a few weak setae. Sternite 5 simple, (1m); BearCanyonBridge, Lociel-BisbeeRoad, 5300', darldypigmentedalongposteriormargin.Epandrium 29.vii,l965; H.B. Leech (If CAS); Yank's Spring, with short, sparse setae dorsally, laterally with 6-7 SycamoreCanyon,TumacacoriMts,4OOO',H.B.Leech lOng bristles Surstylus (li'igs. 41, 43) short setose (1m CAS); 7 mi WNW NOgaIes, Clllabssas Canyon, posterolaterally, posteroventral setae dense, thick, 3850', desert grassland, ix.1972, A Newton(lm1f); 8 and relatively short; some long thin bristles also mi NNW Nogales, Walker Canyon, 3900', oak wood- Vol. 6, No. 3-4, September - December, 1992 173

53 grea r gen elg. Spuler stated that the holotype of B. singularis Figures 50055. Male NorrbOmw,. Left lateral termmaIia; was a male from Kettle Falls, Washington, collected ;t:S; ~~:~::;ctu=;po~terminalia by Melander; whose collection was later sold to the USNM. Wefoundnospecimenlabelledasholotypein the USNM type collection, but In the general collec­ land, dung, ix.1970, A. Newton (lml!). Pima Co., tionthere'\\'8S a maleofN. scripta, originallyfromthe Melander collection, with appropriate locality data. Considering the discrepancies with other USNM Spulertypes (e.g. what are labelled as the holotypes human excrement, J.M. Aldrieh (11 USNM). IILI of Borborus roldtrons and R immen.'la are probably NOIS. Champaign Co., l.vi, 17.v.1925, M.W. only paratypes, andwhatis probably the holotype of Shackleford (lmlf CNC). KANSAS. Manhattan, the formeris18be1ledas a paratypeofanunpublished name, see also comments abouttheB. peltastes types :;3~:;==~~':~~~~~~ under N. lacteipennis), it would not be surprising if Carlsbad, Sitting Bull Falls, 4600', 23 27.'lii.1975, this male was the holotype. We have made this dung, S. Peck (lOmBt). Guadalupe Co., Santa Rosa assumptionasthe specimenmore orlessfits Spuler's Lake SLate Pmk, 3-4.vili.1984, dung, B.V. Brown deSCriPtion, aIthOllghtheI8tterisnotdetailedenough (lm).LincolnCo., lOmiNSielTaBlanca,lQ.26.vi.1979, to eonfirm with eerLainty that B. singularis is N. alpine meadow, 11000', carrion S.&J. Peck (2mlf). scripta. We originally assumed that the name B. ~~~~ Comanche ~ ~ Witta Mis ~~' singulal'is colrespunded to the tpecies we are now calling N. singusta, because of Spuler's statement thaf.f.behind fem.nrisstronglySWOllen TheSize Ofthe reation Area, creek, bottom woods, S.&1. Peck (1m); gena is larger in that species, however; the genal Texas Co., Guymon, 21.vi.1941, L.H. Banker (1m heightismorethan0.5timesthe longdiameterofthe eye, whereas in the Kettle Falls male itis 0.4 times. much closerto Spuler's statementthatthis ratiois l/ (6m5fFSCA); Randall Co., 3420', Palo Duro Canyon 3 ThehindfemnrisonlysJightlySW011enintheKet.tJe StatePark,7-9.vi.1979, mesquiteprairie,S.&1. Peck, Falls male, but in his key, Spuler was comparing it ('7mlli); SanPatricio Co., WelderWildlife Refuge near with that efN. fumipennis (as B. aretieus) which he VernaMills, l.xi.1988, ondungscarab(Canthon sp.), hadnotseen, andwhichMalloch describedashaving J.Heraty(lm1!);Terrence Co., Dryden, IV-V.1954, O. thehindfemur"normalinsize". MostspecimensofN. 174 Insecta Mundi

scripta we have seenhave the wing membrane whilr TJWI'OX KaLepisternum bare below level of dorsal ish,whereasSpulersaiditwash.yalin.e,withtheveins setae;anepisternumwithnonpruinose arealarge,its brown, but the Kettle Falls male IS one of the SPec!­ dorsal m.argm approxnnately level Wlth to slightly mensfjttipgthelatterdescription. OtherthanSpuler's dorsaltodorsalmarginofspiracle; pleuronotherwise statement that there were four rows of acrostichal lightly pruinose except narrow bare area on meron; setae,whichdoesnotfit8ll) speeiesof~''101Tbomia,the scutum and scutellum uniformly lightly pruinose, othercharactershe listed, suchasthe hindtibiawith subshiny. Wing membrane traneparent, with light a row oflong thinhairs onthe extensor(dorsal:) side, browntint; all veins pale brown; secondcostal sector the location ofr-m and the ratio ofthe sections ofthe 20-22X as long as third Midleg dark. brown, tarsi medialveIn, theshortanstalength, andvanouscolor brOWIi; fil'St tarSOmere with slightlyerilargedventral and chaetotaxy characters, all fit the Kettle Falls bristles. Hindleglong-setose, brown; tibiawith. small maIe as well as several other Nearctic Norrbomia anteroapical spur and large apicoventral spur. Hind species. They do not contradict tire idea that tire tibiaofmale WillImany long pOsterodorsalhairsover Kettle Falls male is the B. singularis holotype, but twice as long as tibial width. Hind femur, at least in neither do they confirm this Wlth certainty. The male, strongly swonen, Wlth 2 thin dorsal bristles. surstylus ofthis male differs slightly from Figure 41 Pulv:illi small, narrow, less than halfas longas claw; in that the posterior part projects a bit more as a padS widely separated. Male abdomen: Syntergite ventrallobe andthe setaeontheposterim m8:Igin are 1+2 uniformly pruinose brown, about as long as slightly smaller. We do not consider this variation tergites 3 and 4 together. Membrane ofsegments 3-5 significant and therefore regard B. singularis as a Wlth smallsetae. Stermte 5 Slll1ple, darklypIgmented junior synonym ofN scripta. alongposteriormargin. Epandriumwithsparse,short several eXamIned spemmens were colIeCted on a setaedorsany,lateralIywith4"5IongbriStles.Surstylus S(larab (Canthon sp.), and the white winged flies (Figs. 44, 46)setuloseposterolaterally,postel'Oventral briefly described by Moulton (1880) as riding on a setae forming straightrow of8 stout, straight setae; dungbeetle inMissouti are probably this species. H. anterior part ofsurstylus elongaLe, with small, carl- Howden and J. Martin (pers. comm.) filmed white- nate anteriorly rounded inner ventral lobe. Cercus winged flies riding on Canthon humectUs (Say) In Wlth a few small setulae posteromedially and Durango, Mexico in 1964. Consideringthe location, posteroventrally;apieallywithelongate,truneatelobes these flies probably also were N. scripta. separatedbydeepcleft;cercifuseddorsally.Paramere (Fig. 45) broad near middle. tapered l;o MULe apical Norrbomia singusta, new species process; anterior surface with broadly triangular (Figs. 44-46, 89-93) process near middle and quadrate lobe preapiCaJIy Basiphalluswithlong, naITOW epiphallus. Basalscle Description. Body length 2.0-2.5 mm. Body color rite of distiph81lus undivided, apic81ly with single, reddish bi'Own to black; antenna and legS brown; medial, dorsally projected lobe; distal part of halter white or yellowish. Head: Aristallength less distiphallus larger, lightly sclerotized; anteroventral than internbriss& distance, aristal hairs shorLer surface with strongly serrate lobes flanking base of than aristal width. Frons with medium size stout median process. Female abdomen: All

(Fig. 91) smooth, with few, "Y\<-eak bristles. TeJ:gites 6 and7 undivided;tergite 8 bipartite; epiproct(Fig. 93) simple, bare except for 2 bristles. Sternites 6 and 7 entire, sternite 8 tripartite, with dark, very large posterolateral lObes; medianpari nalTOW ana weakly

Figures 56-78. FemaleNorrbomia. Figs. 56-60, N. frigipennis. 56, abdomen dorsal; 57, abdomenventral; 58, abdomen lateral; 59, spennathecae; 611, ePJpfucl and reici Figs 61-65, N /itld.permis 61, abdOmen dorsal; 62, abdomenlTentral; 63, abdomen lateral; 64, spermathecae; 65, epiproct and rerei. Figs. 66-71, N. {umipennis. 66, abdomen dorsal; 67, abdomenventral; 68, abdomen lateral;69, spennatbecae; '/11, epiproct and cet'ci; '71, hYPoproct, cerci and sCIerite anterior to hypoproct. Fig. 72, N. yukoneTl8is, hypoproct, rerei and sclerite anteriorto hypoproct. Figs. 73-78, N. lacteipennis. 73, abdomen dorsal; 74, abdomen ventral; 75, abdomen lateral; 76, spermathecae; 77, hypoproct andrel'Cl; '78, eplproct and rel'Ci. Vol. 6, No. 3-4, September - December, 1992 175 176 Insecta Mundi sclerotized, steuute 8 large, its postero1atera1lobes Bo7 bonts bilineatus Olimshaw 1901.75. Syntypes extending lateral to abdomen in dry specimens. male female (BMNH). UNITED STATES. HA- Hypoproct emarginate and bare anteriorly, setulose VIAll. Kona. posteriorly Spermatheca(Fig 92) subspherical,flat- Borboros minutusJohnson 1913:449. Holotype male tened at top and bOttom, with large basaI apOdeme; (MCZ). BERMUDA. SpanishPoint, D.vii, Spaeth. neek kmger than main part. Borbonl8brnllisetusMaJloch 1913·365 I eetotypemale [here deSIgnated] (USNM), UNtIED STATES. Wl\SlilNG'I'ON,D C [WithfOl1Owmglllbels'*Nat Mus.,28.4.12,J.R. Malloch"; red"TypeNo. 14948 ynomys urrow,. oc erot. U.S.N.M.",and"Bomon1sbrev.iBetaMalloeh [male symbol] Type"]. Borborus (Borborilkts) marmoratus. Spu1eJ: 1925.11, Paratypes. Same data as holotype (1Om9f CNC, and authors not Becker. USNM). UNITED STATES. ARIZONA. Navajo Co., Borborus (Borb;;rilkts) sordidus. Duda 1923.84. 15 mi SE Holbrook, 14-16.vii.1979, 5300', grassland Coprom;yza (Borborillus) sordidn. Richards 1930.313, carrion traps, S.&J. Peck (1m). 1965:720; Hackman 1977:399. Norrbomia sordida: Papp 1988:394. Distribution. Known only frOm Saskatchewan and A • ..ll"lzon&. DescriPtion. BOdY length 2.5"3.0 mm. BOdy, legs andantennae darkbrown to black, heavily pruinose Comments. Norrbomia singusta is a distinctive exceptshiny areas onpleuron, gena, terminalia, and anteriormarginsoftergites;halterpalebrown.Head; 8Cnptaan.d~mexuxma, 0 w s n ve Aristallengthlessthanintervibrissaldistance;aristal =~~ch~~~=e(row~sUl'Btylar es an em e erse an nor groove on hairs shorterthan aristal Width. FrOns entirely pru' inuse, interfrontalandorbitalplatesgoldenpruinose, sternites 6 and 7) Within this group, N. singusta is distinctive for Its undiVIded fernaIe sternite 4 and Its inter.oeningareasandbaekofheaddarirer.Interfrontal characteristiclongmalecerci. IthasbeenooUeetedin bristles in 5 small pairs; 6-7 inclinate inner orbital mammal burrows, butotherwisenothingis knownof setulae. Face uniformlybrown, pruinose. Eye height its biology. The specific epithetis a coinedword and en. 2.5X genalheight. Gena(Fig. 19)shinyblack,Witb is considered a noun in apposition. broad, whitish, densely pruinuse ventral area (ex- tl'eme ventral margin very n8l'l'OVl.'1.y nonpnliriOBe); Norrbomia sordida (Zetterstedt) nonmicrotrichuse area approximatelyhalfaswide as (Figs. 9,13, 19,25,47-49,94-98) pruinuseareaatnmrowestpoint,broadetungslighUy at anterior margin. Thorax: Katepisternum nonpruinosebelowlevelofdorsalsetae;anepistemum Cop::t'Th80~Ze~dt 1847:2484. Leetotype prUiIiOSe except for tinY isolated bare area iIllDlEldi- ately anteroventral to spiracle, and often very small soi'dida Svec. unik." in Zetterstedt's writing]. bare areainantel'O'V"entral corner; pleuron,including Copromyza sord-ida var. nigriteUa Zettersiedt meron,otherwiseentirelypruinuse. Scutumandscute· Hum heavily pruinose; pruinosity brown between acrosticlial bristles, Silvery between afu'OSticlials and E erod with onI tin blue reetan Ie which dorsocentrals, brown along dorsocentral rows, then ass rO e oc y ersson, , pel'S comm ), and a label with *C nigritelJa (male prescutellarpair of dorsocentrals large and well dif· symbol] anvar. sordid.?"m Zetterstedt'swntirig]. ferentiated (8 pairs are well developed in other spe­ cies) Wingmembrane transparent) with light.brown

Figures 79-98. Female Norrbomia. Figs. 79-83, N. mexicana. 79, abdomen dorsal; SO, abdomen ventral; 81, abdomen lateral; 82, spermathecae; 83, epiproct and cerci. Figs. 84 88, ~7\.l scripta. 84, abdomen dOl'Sal; 85, abdomen -..entral; 86, abdomen lateral; 87, spermathecae; 88, epiproct and cerci. Figs. 89-93, N. singusta. 89, abdomen dorsal; 90, abdomen "entral; 91, abdomen lateral; 92, spennathecae; 93, epiproct and cerci. Figs. 94 98, .\T. sardida. 94, abdomen dorsal; 95, abdomen ventral; 96, abdomen lateral; 97, spermathecae; 98, epiproct and cerci. Vol. 6, No. 3-4, September· December, 1992 177 178 Insecta Mundi tint;allveinspalebrown;secondcostalsector2.0-2.9X New Brunswick, Newfoundland, Nova Scotia, as long as third. Midleg brown; first tarsomere with Saskatchewan IJNlTED STATFS Alabama, Alaska, uniform ventriil bnstles. Hilid1eg very short-setose, AriZOna, Califorma, COlorado, Flonda, Geargia, Hawan, brown; tibia with. single, large apicovoo.tral spur; Idaho,lUinois,Indiana,Iowa,Kansas,Maryland,Massa tibialsetaelessthanhaHaslongastibialwidth, setae chusetts,Michigan,Minnesota,Missouri,Montana,New alongpostewdoIsaledgeofmaletibianotenlargedas Hampshire,NewJersey,NewMexico, NewYak,North in most other species. Hind femur with 1 thin dorsal Carolina. Ohio, Oregon, PellIlSY1vania, Texas, Utah, bristle. Pulvillismall,narrow,lessthanhalfaslongas VJrglllla, Washington, Washibgton, D.C., W'lSCOIlSlll, claw; pads widely separated Male abdomen' Wyoming BERM!IDA MEXICO BajaCaliforniaNorte, Syntergite1+2 pruinosebrownexceptformushl'OOm­ Guerrero,Veracruz. GUATEMALA: AltaVera Paz.EN­ shaped pale median area on wgite 1; synteIgite GLAND. GERMANY. GREECE. SPAIN. Specimens slightly shorterthantergites 3 and4 together. Mem­ collected v-viii, most by sweeping but some taken from brane of segments 3-0 bare or With a few minute barsedung. setulae Sternite 5 simple, darkly pigmented and broadly bilobed posteromediaIly, setae restricted to DistributIOn. A common Holareti.c species, occurrmg posterior half and middle region. Epandrium with south to Guatemala in the Neotropiea1 Region. Papp sparse, medium-length setae only. Surstylus (Figs. (1988) suggests that Afrotropical records are 47, 49) setulose posterolatera1ly; anterior part of misiden:tifications, and records from India aJackman surstyluselongate,withlarge,carinate,innerventral 1977) are unconfumed. Although N. 80rdidais reliably lobe; -inner ventral surface of surstylus WIth 2 very reoorded from Hawan (Tenono 1968), an other Pacific large bristles extendingbeyond ventral lobe Cercus indioo group specimens we examiIled belong to other with a few small setae posteromedially and species. WebaVe seennospecimensofN. 8OrcJi(Jaorany posteIo..-entrally;apicallywithelongate,broadl,eari other NorTbomia from the Neotropical Region south of nate lobes separated by deep cleft; cerci fused dor­ Guatemala. sally. I'aramere (Fig 48) broad nearmiddle, tapered to narrow, truncate process; anterior surface with Comments.Norrbomia 80rdidais a distinctive species, broadly trIangular process lobe preapicaIIy. differing greatly from its New WOrld congeners, from Basiphalluswithlong, narrowepiphallus. Basalsele whichitismesteasilyseparatedbyitsdense pnDnosity, rite of distiphallus with sheath-like lateral lobes, theverynarrow,anteriorlybroadened,nonmicrotrlchose medial part slightly bilobed apically; distal part of areaonthegena:,thestripednotwn;andmanyabdominal distiphallus with loop-like distal lobes; ventral pro­ characters. This species is synanthropic and copropha­ cess With narrow process projeCted proXimally be- gous, anaIWl pro6abI.y attamedits Wide diStributiOn by tween ventral basal lobes. Female abdomen: moving with man from the Old Worid. 'The elosest Preabdominalscleritesundivided, darklypigmented; relatives to N. sordida are OldWorldspecies. pleura(Fig.96)weakly slJ:iaLe, withfew,smallsetulae. Tergites 6 and 7 (Fig. 94) emarginate posteriorly; Norrbomia triglabra, new species tergite 8 emarginate anteriOrly and poStei"ioi'ly, pos- (Figs. 50-52) terolateralCOnIers detached; epiproct(Fig. 98) short, setUlose, with 2 large briStles and 4 5 small bristles. Description. Body length 3.8-3.9 mm. BOdy color Stemites 6 and 7 (Fig. 95) posteriorly emmg.inate, darkreddishblownto black; antenna, basesoftibiae posterolateralCOnIers ofsternite7 detached. Sternite and tarsi luteo~terstem bro~knob whitish. 8 bipartite, eachhalfstronglytapered anteriorly,less strongly tapered and very dark posteriorly. Hypoproct setUkJse except for namJ\1\' anterol8.ter8l lObeS: sper. matheca (Fig. 97) teardrop-shaped, with slight con.stric- lateral to oeelli and small nonmierotriehose shiny tionwhere itbegins totaperto neck, whichis en. 2.5X as areainfront ofanterior ocellus. Interfrontal bristles longas mainpart; large apical apodeme present; butno in5-6 pairs; 9·10 inclinaLeinnerorbitalsetulae. Each basal apodeme. side onace dark except along carinate middle partof face. Eye heIght roo 2.OX genal heIght. Gena dark Material examined: Lectntypes of C. 80rdida and B. :reddish brown, micrctrichose narrowly anteriorly brevisetus, holotype ofG. nigritella (see synonymy), and andfairlybroadlyventrally(extremeventralmargin over400 specimens fromthe following areas: CANADA narrowly bare), nonmicrotriehose area broad, trun- Alberta, British Columbia, Manitoba, Ontario, Quebec, Vol. 6, No. 3-4, September - December, 1992 179

eale anteriorly. TJWI't/,X KaLepislernum bare below A female ill Lire MCZ may also be Llris species. It level of dorsal setae; anepistemum. with. dorsal mar- bearsthefollowingdata.: GEORGIA. Clayton, 2000- gm of nonnucrotrichose area approXlDlately level 3700 ft, VI. 1909, W.T. DaVIS, ·onCanthon chalettes with ventral margin of spiracle, nonmicrotr.ichose ball". area immediately ventral to spiracle not isolated; pleuronotherwiselightlypruinosej scutumandscutel- lum uniforml heavil ruinose. Win membrane ansparen , Wl g rown n; veIns p e (Figs. 53-55, 72) brown; second costal sector 3 ox as long as third Foreleg setose, brown, tarSI and base of tibIa lute­ ous. Midleg brown, tal'Si and base oftibia luteous. Description. Body length 2.1-3.2 mm. Body color, Hindleg short setose, brown; femur with 1 thin, including antennae, face and legs, black, halter pale erect, dorsal bristle; tibia with small anteroapical brownoryellowish. Head: Aristallength subequalto bristle and lar e curved a icoventral sur' intervibrissal distance; aristal hairs 00. 3X basal pos ro ors I se ae SIml ar 0 er se e, shorter than tibial width Pulvilli small, widely areas beside ocelli, if present, smaller than ocellus. separated, together much smaller than tarsomere 5, covering only basal halfofclaw. Male abdomen. Syntergite 1+2 uniformly pruinose brown. Mem­ brane of segments 3-5 with 3-4 small setae per Katepistemum usually bare below level of dorsal Slriij)le, ar y plgmen ong po nor margIn. setae, anteroventl'81marginsometimesmicrotriehose; Epandrium with dense, short setae dorsally, later anepistemum with tiny, isolated nonnrlcrotrichose allywith4-5longbristles, dorsolateralbristlelong­ area immediately anteroventral to spiracle, nmin est.Surstylus(Figs. 50,52)setulose poslerolaterally, nonmicrotrichose area small, its ~ors:u ~=nwell setae relatively short, less than half as long as ventralto levelofventralmargm0 SpIrace;peuron, includingmeron,otherwiseentirelylightlypruinose; surstylus; antenorpartofsurstylusquadrate, Wlth blade-like inner venirallobe. Cercus with a group scutum and scutellum uniformly, heavily pruinose. of7-8 setaeposterodorsally; posteroventrallyshort Wingmembrane transparent, with light brown lint, setose, elongate, apex expanded, elublike; cerci all veins brown; secon~stalsec~.8-3.~ asthird.Forelegblack, ~eessom~esre.= separated by deep apical notch, cerci postemmediallycontigl.lol.lSdorsally Paramere(Fig tibi~k, ~omere with slightly enlarged 51) broad near middle, tapered to acute apical process; anterior surface withtriangular lobe near single, very long apiooventral spur, femur with 2 3 middle. Basiphallus with long, narrow epiphallus. erect dorsal bristles on distal half. Posterodorsal Basal sclerite of distiphallus undivided, apically bristles ofmale hindtibia slightly shorterthan tibial with single, medial, dorsally projected lobe; distal =%:l:;:t:;~if;:::tt'y~~~lt= part of distiphallus larger, with quadrate distal venfrallobe Remote unknOwn Membrane of segments 3 5 bare or with 1 2 minute setulae. Sternite 5 large, dark; posteromedial area Holotype. Male (CNC), UNITED STATES. KEN­ pale and densely nricroselulose; ventralpartofstem- TUCKY. Vv"nitley Co., CumberlandFallsStatePark, ~~h~:e~~e=~:t:f:;:~'~~'~;::': 29-30.vi.1984, dung trap, S.A. Marshall. withsparse, shortsetae dorsally; laterally andpeste Paratype. Same data as holotype (1m GUE). riorly with a few longer bristles. Surstylus (Figs. 53, 55) with large innC! ventJ: al lobe, narrow distally, Distribution. Known only from Kentueky. rounded along outer surface, broaderthan thatofN. famipennis and narrower than that of N. costalis; Comments.This speciesbelongstothe frigipennis :7tl::p::~:=,~:~~~: ::o~ group, and can be most easily distinguished from other species in the group by its distinctive male view. Cereuswithbare posteriorareabelow setulose, cerei. Norrbomia triglabra is named for the three rounded, slightly raised part and above strongly glabrous areas that surround the ocellar triangle. elevated posteroventral part. Cerei divided by nar- 180 Insecta Mundi row, deep, almostparallel-sidedcleft Paramere (Fig. tic region as well as the northwestern Neareti:e are 54)verybroadon distal half, abroptly taperedtosbort required to properly clarify the N oostalis complex apIcal process; 5 setulae alongpostenorsurface and2 =:'i;'::;u7~:i"~ie:;:~:;h~t3 ACknmvledgements vided, apieallywithsingle,medial, dorsalI, projected ::e~~ t~ c~rs.of the mu~s~d~ lobe; distal partof distiphallus larger, with flattened the e 0 section or elrgenerous oans 0 speC!- distal ventral lobe and paired, dorsoventrally flat­ mens. Special thanks are due to Dr. P.I. Persson tened, dist.al dorsal lobes Female abdomen' Gener- (NRS) for locatingthe types ofC. fumipennis. Draw- ally siDillar to N. fUmzpennUJ, but WIth considerable ings of the female terminalia are by I.P. Smith. P. variation in development of selerites of stemite 4, Malikul assisted with the SEM photographs B V rangingfrom4 heavilysclerotizedsectionscharacter­ BrOwn, T.A. Wheeler, G.C. Steysklil, RE. White and isticofN. fUmivennis andN. 008talisto2 SIDa)] medial R.J. Gagne kindly reviewed the manuscript. sclerites,with outerpairreduced Sternite8hipartite, mnerpostenorcornersofeachhaIfralsedmto strOng References c~ Hypoproct deeply emarginate anteriorly, DudlG O.W. 1923. Revision der altweUl:ichen Arlen with . d scleriteinemargination, this sclerite sepa­ derGattungBorboros(Cypsela)Meigen(Diptera). rate from hypopr'OCt and deeply cleft such that each Archiv fiir NaturgeSChichte 89'35-112 arm is as long as median length of hypoproct. Farris,J.S. 1988. Hennig86 Reference, Version 1.5. Holotype. Male (CNC), CANADA YUKON North- emRichardSOn Mountains, ErebiaCreekbasecamp, Grimshaw, P.R. 1901. Diptera. FaunaHawaiiensis 67°68'N 186°29'W, dung cups in wet tundra, 1 3'1-77 8.vii.1987, SA Marshall. Hackman, w. 1977. Family Sphaeroeeridae Paratypes. Same data as holotype (3mlf GUE, (Borboridae), pp. 398-406. In M.D. Delfinado and USNM). CANADA. YUKON. locality as abOve but: D.E. Hardy, eds., A Catalog ofthe Diptera ofthe mammal nms near creek (2f)j carrion trap in wei OrientalRegion, Vol. III, SuborderCyclorrhapha tundra (2f)j flood plain ofcreek (2f GUE, USNM). (excludingDivisionAschiza). UniversityPressof Hawaii, HOnolUlu Distribution. Known only from the Yukon. Johnson, C.W. 1918. The dipteran fauna of Ber· ~men~s. Norrbomia yukonensis resembles N. Dluda. AmIals of the Entomological Society of mzpenms, whichalsooccursinnorthwestemNorth America 6:434-452. America, so closely that examination of the male genitalia is necessary to reliably tell them apart. There are several Paleal'cl:ic species nfibe N rostali.s complexthat areverycloselyrelatedtoN. yukonensis speciesdescribedhere differs fl6DlN. eostalisandthe verysimilarN. cryptica (papp)primarilyindetails of Knab, F. 1915. Dipterological miscellany. Proceed­ the sw-s!;ylus. Bothofthelatterspecieshavebroader irigS Of the EritomOIOgiGaI SOCiety Of washington inner lobes and more elongate clusters oflong basal 17:38-40. bristles The Shape Of the scIerite anteriOl' to the hypoproct also differs among these species. The Malloch, J.R. 1913. Descriptions of new species of variability in stemite 4 ofthe female is here consid­ Americanflies ofthefamily Borboridae. Proceed­ eredintrwmecific,butitshouldbenotedthatthemain iDgsOfthe UriitedstatesNation8lMuseum44:361 characterusedbyPappto distinguish the Mongolian 372. species N Crj'Ptica fromN costalisisthereductionof the female fourth stemitetotwo scleritesinthelatter Malloch, J.R. 1915. Two new North American species. As is the ease in many groups, taxonomie Diptera. Bulletin of the Brooklyn Entomological studyofadeguatecollectionsfromtheeastemPalearc- Society 10·64-66 Vol. 6, No. 3-4, September - December, 1992 181

Marshall, B.A 1987. Systematics ofBithecn, a new Richards, O.W. 1967. Family Sphaeroceridae genus ofNew World Sphaeroceridae. Systematic (Borboridae).ln A.E. Vanzolini and N Papavem, Entomology 12:355-380. eds., A catalogue of the Diptera of the Amencas south of the United States. Departamento de McAlpine,J.F. 1981. Morphology andTerminology Zoologia, Secretaria da Agricultura, Sao Paulo. - Adults, pp. 9-63. In McAlpine, J.F. et aI., eds., Fasc. 72.1-28. Manual ofNearctic Diptera. Volume 1. Research Branch, Agriculture Canada, Monograph 27. Sivinski,J. 1983. The natural history of a phoretic sphaerocerid Diptera fauna Erological Entomol- MouRon,J.T. 1880. F'fies nmngon a 'rumble-dung. ogy 8:419-426. The American Entomologist 3:226. Sivinski, J. 1984. Sexual conflict and choice in a Norrbom,A.LandK.C. Kim. 1985.Taxonomyand phoretic fly, Borborillus (rigipennis (Sphaero- phylogenetic relationships of Copromyza Fallen ceridae). Annals of the Entomological Society of (s.s) (Diptera: Sphaerocendae). Annals of the Amenca 77:232-235. Entomological Society ofAmerica 78'331-347 Spuler,A. 1925. NorlhADlericanspeciesofBorbOTUS Norrbom, A.L and S.A. Marshall. 1988. New MeigenandSro,t<>phoraRobineau Desroidy.Bu!- species and phylogenetic analysis of Lotophila letinofthe BrooklynEntomological Society20:1­ Lioy (Diptera: Sphaeroceridae). Proceedings of 16. the Entomological Society ofOntario 119·17-33 Stenhariimar,C.I855.Sk8iidiri.aVlensCOpromyzmae Papp,L 1988. A:reviewoftheP..frotropical speciesof granskadeooh beskr..fne. Kongoliga "Vetenskaps Norrbomia Gen. N. (Diptera: Sphaeroceridae, -Akad.emiensHandlinger(1853)74:258-442. [year Copromyzini). Acta Zoologiea IIungarica 34:393- usually given as 1854, buL not published until 408. 1855 (p.I. Persson, pers. comm.)].

PaPPIL.andA.L. Norrbom.1992.Taxonomyofthe Steyskal, G.C. 1971. Notes on some species of the Paleotropical genus Achaetothorax Hedicke genus Copromyza, subgenus Borborillus. Jour­ (Diptera: Sphaeroceridae). Acta Zoologica nalofThe Kansas Entomological Society 44.476- Hungarica 38:321-344. 479.

Richards, D.W. 1930. The British species of Tenorio, J.A. 1968 Taxonomic and biological stud- Sphaerocerietae (BOrbOridae, IJiptera). PrOCeed­ ies of Hawaiian Sphaerocerietae (IJiptera): Pro" ings of the Zoological Society of London. 18:261 reedings of the Hawaiian Entomologieal Society 345. 20:169-212.

Richards, O.W. 1965. Family Sphaeroceridae. pp. Zetterstedt, J.W. 1847. Diptera scandinaviae. 71&726 In A Stone et ai, eds, A catalOg Of DispoSita et desCriPta. (part)6:2163-2580. LUnd. Dipteraof.AmericanorthofMexioo. U.S. Depart mentofAgriculture, Agricultural Handbook No. 276. 182 Insecta Mundi