The North American Subspecies of Culicoides Variipennis (Diptera, Heleidae) L 7

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11111 1.25 1111,1.4 ""11.6 111111.25 11111 1.4 11111 1.6

. ,

MICROCOPY RESOLUTION TEST CHART MICROCOPY RESOLUTION TEST CHART NATIONAL BUREAU Of STANDARDS-J963-A NATIONAL BUREAU OF STANDARDS-J963-A REFERENGE t~O'F LOAN I 00 • \ The North American Subspecies of Culicoides, variipennis 1

\ (Diptera, Heleidae)

?'!. ~ ~. ,~ ~ r- ~ o LO ~ ~ 1= ~ f'- ~ o ...... (J) ~ <....) ~ r~' u....I ~ - Q en ~ ~ by Willis W. Wirth and Robert H. Jones • \

Technical Bulletin 1170 • UNITED STATES DEPARTMENT OF AGRICULTURE ;/

•

ACKNOWLEDGMENTS

\,-(\ are gn'atlr illllt of Agl"iellitllrf> ('.\Xl, fOJ: !'lIgg('sting thf> taxollomic.' study ,)f the "uria!:iOIl ill II0I)llla t iOlls of I ht, ro rii /U'IIII i.q l'ompll'x, a lid for ('a \lill;: ('ertnin USt'fui ('haract:(>rs to our at-t('lItioll; allli to tht'lIl, as well as tlw flll\owin!!, 1)(>rson;; and .institution", ror tht' loall 01" Jlr('';PlIlaliriul whif'h mude this siu tpxt· reeol"lls as indicated in 1):1r(>nth(',,(''') : ~In:;. Eli7.alwth ('. lW('k, Floridn ~tat:c Boanl of Health, JaeksOlwil1<.' (FL,\): {:. D, Hllt'l('r, l'lli\'l~)'''ity of ..\l'i7.()lIn, 'I'm'soll; lI, Dietrieh, Corn(>l1 lJni"l'l'"Hr, Ithaea, ~. y, {( 'on) : D, K Ilowell. Oklahoma ,\, and l\£. ('(1l\e!!'I', St:il1\\':lll'r (OKLA): ~1. '1'. :rall1l''', '1\'aHhiugl'!I1 ~t:lt\, Collt~g(', l'nlhnan ('IYASll): ll, n, Pratt, COlli 1II1lllkn bl(' nisl'US(> ('('111:1'1', t', ~, Publie Health Sen'it-p, "\I"lallla, <::1., 1('1)('): L. W. tinnil'. \'Ilh'('r"iiy of ::$>brn~k", Lineolll (NEBR); (', K nkhnr!\";, I'. ~. 1'11hli" 1I('alih ~<'I'\'ke, :l'P111])(> , Al'i7..; .II. II. Hoss, Illinoi,:; ::\:lil\l':tl lIi:;tol'Y ~Ul'\'('r, lJl'lHlna (lLL); R. E. Ryek.lliln, Rehool of Tropirnl atHl 1'1'!'\'\'nf'in' ~I('dieitl\'. Lonl:1 LitHia, ('alif,: '1Y. E. ~nn\\', T('nnt'i'i'(>(> Valler ,\ut\lOrity, Yfil"nn Dalll.•\Iah:un:l 1'1'\'.$; I'a\ll K ~pan!!l('r, \-Ilin'r:;ity of ~li"~()lIl'i, (',dlllllhia: and '1'. O. 'l'linh'Ill'I', ('Olol':llln .\. &:\1. ('011 I';!l' , 1."01'(: Collins (COLO), (".lip,,;; :;i:ltl'(1 othc'l'\\'i,,(', Iht' Illntpl'ial i;; fl'olll tlil' U. R. :-\ation:ll ~II1i;(,UIll ('ol1('C'tioll ill \\'tlHhillgtllll. D. C. (i'or t h(' 10:ln or dOllntion of "11l'dllH'n~ 01' r('lat('() I:'pel'il':; 01' Cllli('oi

• •

Contents

l'ugc Introduction.,•.. Taxonomic relation,: ___ . Taxonomic differentiation____ ..___ ...... _.... 5 C'lIlicoides t'ariipennis mriipennis (Coquillett) ___ . _. _ ...... ] 2 Culicoides ('ariipennis llll.~tratjs 'Wirth :1nd Jones, new subspecic~_. I.) ClIlicoides t·ariipenni.~ alberten.~is \rirth and Jones, 11(,\\" "llb~pecies____ 17 C'lllicoide$ rariipennis ,s()norell.~i.~ Wirth and ,Jon~s, new sub"pecie~._ IS C'wicoides t·(lriiJ!enni.~ orridenlali.~Wirth and .Jones, up,," ,.;ub::;pecies___ 21 Statistical eyaluation of fOUi' charnct<'r:' of th(' ."ub,:pcC'i(';< of the t'arii"Pnni,~ complex.. ---..... 28 General consideration:,_ -. 32 Refer('nce~ - .•-- .• --- :34

Washington, .D. C. Issuc(l October 1037

For sale by tho Superintendent of DO('I1I11(,l1t;;, l". ~. (;O\'ernm!.'lJ[ PI illting Ollie!.', • Washington 25, D. (' '- Price 1.5 cents III • \ \/ The North American SUbspecies of Culicoides variipennis (Diptera, Heleidae) L 7

\ by Willis W. Wirth and Robert H. Jones

i';lItoHHI/"f}!1 R("~('''I'('II .f);'l'il

Introduction

It has been kn(j)\"11 1'01' :;OUle tillle that w('sterll spc~illlen::; of Cuii caides 1'(U'iipelllli8 (Coqllilletl') art' smaller and palcr: with brighter wil1rr mal'kinrr::; than tho::e in thp Ea::;t. In 1\J5~ the senior author, ill his study (rf the lIeleidac of C;difomia, was unable to analyze the differences with the materi:ll then Onhlll1cl at the r. ~, ~lltionlll J(useum, largely berllnse :-;om(' CaliJ'ol'l1in. :::pccinH.'lls snperiicially l't'sembled those from the Eastern ~t,ltes more close 1v th,lIl other west ern incliddullls. In the last fr'" wars 8e\'e1'a1 tllx6'nomie ·workers in (}ulicoides have called attention te') tL number of structural characters which are useful in dilferPlltiatinu some closely related species. These ('harneter::; btlYl' lJeen applied tU1{tlH~'Y OlH'S adcled ill the present study. Culicoide8l'(lI"iipennis has been demonstrated to be a IJossible yector of the bluetongue yjI'us di:,ease of ::iheep in Texas (Pri<:e and Hardy, lU;){) , Large lllllHber::; of thi" :-pel'iC':3 1u1"e been identifietl from material collected in slIrveyS' 11l,11le in connection with bluetongue out breaks in the Southwe"t ("Tirth n ltd Bottimer, 1!)5G). These develop ments have pro\'ided both the inet'ntiw and the material to undertake a detailed taxonomic mutly::;is of th(' mriation in the variipennis com plex, Our results arc publish('d llO\\" to make llalJleS n.ntiln.ble for the major sllbspeciiic population::;, and to point out some indications or biolorrical differences which Illay assume importance in disea::;e trans mission and contro1. .

TAXONOMIC RELATIONS C1llicoides var'iipennis l Coquillett) 1901 belongs to the sllbrrenus MonoclLZicoides which Khalaf (195:2) proposed, with nllbecldosus (Meigen) 1818 as type. Kh::Llnf included fuUl' groups in this sub genns, of which only one, the mlbeculos!ls group, is concerned in the present problem. Seyen species were included by Khalaf in this group: ltegneli Causey 1938 from Thailand, nubeclllo8uS (:Meigen) • from Europe and .Asia, parl'oti Kieffer 1922 from England and the 1 2 'J'EcnXIC'AIJ BULLETIS 11 TO, r, S. DEPT, OF AGRICULTURE

.\Iedit{,l'l'ltlll':L1l al'en 01' Europe and Africa, pUllctieolli8 B{'('ker 1DO:Z :frOI11 the Ml'dih'l'l'HIH'nn al'pa, /'irl/ti Kil'f!'l'1' Ull-" fl'oll1 _Eul'l)pe~ stigma PJrigpn) tHIS 1'1'0111 EIII'0lW nnd XOI'tlt ,\/'I'ic'a, all(ll'III'i;J1(,IlIlJ,~ ('0- • qllil1{'(t) 1'1'0111 Xol'flt .\I1\Pl'i('a, 111 addition to tltt' SlH'{'iPS nallll'd 11," Khalaf, we would. assign til(' following 10 till' IIltlJl'(',t/os/(s gl'OIlP: corllulus de Mpillol1 10:ri fl'oi11 ;:';ollth Afl'ica, del/meadi Cause)' 1n:38 fl'om Thrtilnnd, ,qiga8 Hoot and 1[ottmal1 HJ:ri fl'o/ll Wl'~t(,I'Jl Cannd::t~ and llOIIIOt07Jlll.~ Ki('f!,pr 10:21 fl'Olll FormOSlt and ,Japan, Th(' typ(' of' lupneJ:i ('auH',)' 'was ('x::tmilwd :IIHl, on the basis of thp Pl'l'S(,I1('(, of ;;l'IlS01'Ia Oil nlltpllllal sl'gllll'lli-s XI-X'", :tIl O\':t! 110npt'I'fol'ah'd 5(>('1' I1wthpc'a, ant:pllnal ra('io of 1..-1" and aNleaglls with simpl<', llonhifid apex, this specie!'; is l'xcllldl'cl from tlll' IIl1bc('u/08U8 grOllp, The primal'Y ('harnet-ers of the nubN'ulo8ll8 group al'c foun(l in the genitnlin; in the fell1nle tlll'l'P is only 011(' spel'mathN'a, this 11101.'(' Ol' le;.;s elongate, I1sllnll~' ·wit-h hyaline lwr:f'ol'at'ions, Hlldfl'l'qllently alTuntt' :in the mall' thr pal'allll'l'pS art' JlIs('cl basally, tlte Hedl'agus is hifid at. the tip, i'lU\vPlltl'a.lr()ot' of tl1l' ba~ist'yIl' is shOl'! anll simph.., Ihl' dOl'sal I'oot long and Illodpl'atply ;.;lpll<1pl', and till' api('olatpl'al PI'Ol'p;-i ;;e5 of the nillth (('I'gllnl al'p well dp\·elopl'(l. _\dditiollal ('\tal'aeiers :ll'l' a;.; follows: Tht' {'ol1lhinl'd Il'ngth of the last tin' tlnt('lln:tl sl'gm('nts i:, I(';-is thnn that of nH' PI,(,(,t'dillg eight: antennnl s('Il!,;OI'ia :u'e l)l'espni', at 1l1O;';i. Oil seg'nwllts thl'l'l' 1'0 i:('n: a pail' of (Iist-illC'/ fl'ontal tnbel'l'h's al'(' pl'l'~('llt lwt:\\'(,l'lt tIlP fl'olltal carilla nll{1 (hE' IIlesal mal'gins of tIlp lil';;t allt-('1lI1:l1 S('gIllPIl/S ill till' fplllnle; tit" sizl' i" usually I:n'gpl' nwn :\\'I'I'ag'p for til(' !!.'pnl1s: :lnd tftl' l11('sonohlll1 IllIs nll1lH'I'OIl;'; Snlnn dark (>111)('tll1'('S on n li'!!htt1I' baekp.TOII1Hl (('s(,l'pl'in slir/lll1t allll jlw'l'oli) , In (ll'del' to ;.;IlO\Y tht' nahll'p of tll(' s(l'lldul'al ('h:ll'netr]'s lI:-:ed. in til(' (lifi\'l'f'ntintiol1 or \\,plI-n'('(lgnizp

'['AIlL"; I, ('olllpal'isOIl 0/ structural ('hal'((cto'S of 8])(,1'i('8 of Ihl' Culi ('oiel('s l1ul}('('ulostlS {/I'Ollp

Third ;\In.ndibll' AtljPtln!ll ""g- Tibial palpal [(,Plh ment,; wi[ h ,;pinf~" S('gllll'lIl· I:'cn~oritl

Lf/lf/lh' lI'id/h .\'Illllber XUllIlli I" XU1IIlhl' 1'/11"/11(111'< .• 8, 0 11-.15 :3, R-IO ·Hi rtf'11 tit((u/i __ 3.3-3, fi 16 , 3,8-10 () {/ifJ(ts~ 2.5-2. !l 13-14 :),8-10 (i hOlllolomll,' . 3. 0 13-14 :3, 8-10 Ii It lliJCl'lIio,'tI.< • 2, 8-:3, 7 12-1-1 :l,8-IO i 1\-7 /l1,l/dil'olli.< 2,8-:3.0 12 3,8-10 , 5 tie/hi 2, G 10-12 3, R-IO (j -·/iOllla. .)_. f)- 13-J·t 3, 8-10 l'(triilICl1l1i,~ ('()Ill jl\(-X_ I 2, I-a, 0 II-Iii , '3, (1-7) 8-10 ! G-7 • 'flIt; XORTH A~mlU('AX HC13!'5PE('JE:-; OF CTLl('OLDES VARUPEXNIS 3

1. C'u lie0 irl('8 ('0I'll'll t lI,~ de ~[ei lIon, UN'ol'lll'(1 fl'Olll Zullllallll a n<1 Transnutl, South .\fl'i('a. lksC'riptioll and lig\ll'Ni in 1<'i(>dlr]' (I!):)l). Slide's ('xHlllinrH!!lIs with distal bl'n1lches shol-1: with simple I;oints; spinrs presPllt ()\1 n'ntml Slll'f[l('(' of mail! body proximad of the fork. '·J.,arnll' bl'l>l'ding in a!l\lndaIH'(' iIt sem i-I iq II id manlire 01' in grollnd rich ill ol'gani(' ll1:lttt'r tn'Olll1( I drinking troughs and at: th(> edgt':'l of watpr [l1l(ldll's:' (('1t l'lollgat(' :llId ",lig'lIlly Iwnt; thint pnlpa! Sl'g-Il1~'llt ;~::~-:Ui till~(,s as.l(~llg al5 bl'Oa~l, with .::11 a110 \\', t\'ipll' pit: IllHmbble With 1(j t('rtlt; lund tibIal ('olllb \\,Ith Ii :-Pllll'S: SP 11:-0 1'1<\, PI'PS Pili 011 :lIItPIlI1:t1 Sl',p:'(,I'il'lioll HII(lli!!IIl'PS by Hoot alld, f[oll'llHlIl ( W:IT). Slid('s ('XUlllill('(I: 1 fl'nudp, La(' (lpaglls with silll'S of main bod.\' 11101'(, bl'fHl(lh' seh'rotizl'll. tllP HI'Ill:' of tltl' (listal fork StOLItPl' and itll' Yentl'al sUl'fn('e spinos(' as in I'Ill'iiji(,lIl/is S()lIol'('ll",is. L:ll'\':Ll' \\'~'l'L' fOllll<1 (l\\'ata l!);\;;. I'P[l()I'tl'(l as ().~llh'Ii.,·ix) i 1\ [1 Sllln 11 pool of fresh water. :i. Culi('oir/('X illlbf'dt10HII.... (~rt'igell). Hl'(,OI'Lh'd from Em·opl'. He (lescrib('(l and figlll'('(1 b\- Ed\n\l'(ls (l!);~!l). Slides rXl1lllilll'd: ·1: l11al,,':,. (j fplllale::, Chi(k'()ck, ] )Ol'Sl't, En!!l:tnd. ~;~ .Julie J!)·W ••r. .\. Downes. Spel'mnt!lpc:1 pyriforlll ami abniptly bpnt nenl' the l'ntl.'UilCe to tIt<: duct; third pal pal ::;eglnent ~,s-:LT time:" as .long as brOtH!' and pit :small. to medillm-::iizecl: mandible with U-H teeth: hinel tibial comb ,yith (i-T spines: sPt1soria pl'esent on antennal sef,rlnents ITI. YIn-x. ~Iale ,!!l'l1italin, with ninth sternUlll dpeply emarginate; ninth tergunl with :-;1l'LHler apicoiateml processes: basistyle stout on basal half and spinose Olt Illesal margin; nedeaglls with stout expnndl'd, blade-lib' • apices, without \'entm1 spinp;.1; pUl'tllllel'eS with short. pointNlapiel's. 4 TECHSICAL BULLI~'rjX 11 i'D, r:. S. DEP'l'. OF AGHLCULl'{lHg

Edwards (lD39) gaTe the habitat as ]j(luid fl'olll farrnyanl llHUllll'P heaps and green sl ime l'ieh in ol'ganie Illattcr, J)O\yIWS (l!l:,)O, 1D;')i)) ga,ve debtiled notes on till' I iff' h istol'yi 11 I~I1!l'lalld, 6, 01tlicoides plln('tir'ollis Bec1n'r, HN'{)l'ded from the Mediter ranean area, Hedescl'ibed and figured hy Edwards (1!l3!)). SlidE'" examined: 1 female, Tllnisia, ..:\.ft.ieH, j()OG-07, F. S:lntchi, eolL; :2 • lemaJes, Struma ValleT: }[ace1'1'111. ::-icoiland, ;W }(ft." U);}O,.f. _\. Call1pbell ; 1 Illall', GOt'(1on: Hpl'wick, S('otl:Jll(l, :21 ,June l!);jJ; all by courtesy of the A(ol'edull Institute, Edinburgh. Spermathe<:a short and oval, with n. slel1(1cr, thumblike distnllobe; third paJpal segllH'nt 2.6 times as long as broad: w:th n.lllrge, il.'l'l'gulal', ~h:l1l0\\', pitte(L area; mandible with 1:\-1,1, {p('f'b; sl'lIHol'in jll'PHI'IIt on antennal :;e!l'lllents HI, nIl-X; fourth tal'somere distillC'tly cord iJorm. Male genital in. "'ith ninth sternum \'ery deeply ('marginate, the posterior llH'mbrane spicnlate; Jlinth tergum ,y.ith slcndcr ftpieohtpl'al processes llnd a submedian pail' of bluntly rOUJHled caudn l lobes nearly as long; aedeagus ",ith slender hasnl nTIIlS and \'Cry i-ilcndcr disULl points, the main body not bearing yentral spines; parameres ",ith long: pointt'd apicl's. IIabitat among floating green algae at edge 01' small pond (Edw;tl'ds, 1939). 9. O~tlicoide8 '~'aJ'iipenni8 (CoquiJIett). Hecol'ded 1'1'0111 North .America to Mexico, South and Central .American records being ill elTOr for W'llbfle Fox alld Iloll'lllan. ~ee page 12 for diagnosis and dis tribution, Characters diagnostic of this species twe included in n. more complete descJ'iption of I'(ll'ii;Wlllli8 Nll'iipen71i8 below. Sec tables 1 nnd Q 1'01' a comparison o't chantdel's, • THE XOHTH A~[EHrCAX Sl:JBSPr:CIES OF CGLTCOmr:s VARLIPEXNIS 5 TAXONOMIC DIFFERENTIATION In the study of variations within and between species; large samples from many ,yell-distributed localities are highly desirable, if not • absolutely neeessary. 1Ve examined a toULl of about 2,;")00 specimens, of which 1,072 were disseeted and monnt('d on slides; this material came from ;3+ Htatl'S, from !3 Pro"inces of Canada, and from. 5 States in :Mexico. In our search for t:1XOl\Olllie charadel's that might In'o v(' useful in delimiting populations of til(' 1·(lJ·iip(JJlJ1i.~ complex. wc relied primarily on the extensive ('ollect·ion of the 1'. S. National Museum. I\Ye nlso mncle inquil'ies of nll institutions and persons in North Amer ien, who we helie,oecl could furnish us with material from difrerent gl'()£!l'nphic loealities. Those who responded (set' acknowlE'clgments) arc' in Jargl' putt n'spollsiblE' for the nbnndant trlxonomic matel',ial \\'hi('h E'nnbl(>d liS to plot: g'pogmphicftlly tlw (listribution of eharacters in ('ollsidernblp dl'tail. ~\. mnp sh(l\ring the geog'l'aphieal distribution of tIll' S11iJsI)l'('ips is ginn a" fig-m'E' I.

.. ~ 1 .' .j , ~

.' ~\ \

\ " , \ r \ p\ "\~/.... __.::..::.- \

1·'!CWHr; ] .... ·DistriilutiOll of the fi\'(' !-Hlh~p('('i('s of the (·lIli(·oi(I('.~ 1'1I"iifiCl/lli.~ {'Olll pll'X in .'\orth AIllC'ril'lI.

1Ye found the most o\),oious ('olot' difrerences to lw rather nlriable within populations, and sOl11e of the more reliable feutur('s of colora tion were hard to tlefine. After a preliminary triuJ-and-('ITor sifting of dozens of structuml characters, we found foul' fP1Wlh~ characters which, alone 01' in nn'ious combinations, could be used reliably to eharaeterize the major populations. Th('se eharllcters ar(> (1) third paJpaJ segment, the ratio of the length d i "icled b.,' the greatest • hreadth; (2) wing length, in millimeters measuring from the basal ..J~0171--57-2 6 T8CHsrCAL BCLLETIX 1170, e, :::., DEPT, OF ACHfCCLTCRE

arculus to the wing tip; (3) antenna) sensoria, number of antennal segments ill execss of the nOl'mal 11l1l1lbel' (segments nT, nIl IX, X) 'which bear distnJ pits ringed with tufts of short setulae, foul' being Uw highest number possible of extra sensoria (segments IT. Ln. tmel YII) : (.:f) mandible, number of teeth at til(> np<:'x, 1'hes(' ch:.lI'actl'!'1': nl'(' • eonsidel'PCl sllflicil'ntly important. to justify a f::iblllntion of them for e.,,:'h subspe('il's hy locnlity and n dl'tnill'd analysis o'f thl'i!' ":ll'i ability \Yithin and he>('\Yl'en suhspl'eirs, ~\lso wry important al'(' thl' shape of thl' female spermatlll'ea :111(1 thl' pl'esenee 01' absence of linl' spine'S on tIll' "l'ntl':t! SIII'I':ll'(' of tllP sllbapical portion of tIl(' mak aceleagus, Additional lIseful ('haraC'ie'I's ,Yhich arc giyen in our sllh species diagnos(,:-1 inelu pall' and clark ,ying mtll'ki ngs a nd of' the clark-brown Inmetul'e's and 1'a Il' browll infuseai:ion of the 111(>S(JJlotnm. the extl'nt 01' niP s('uh:'lInl' mtll'k illgs, and the ('0101' of' the halter knoh', \~al'iati()n, ill tIll' sb:LJlP of till' Sp('l'I1l:lthl'l'a has IJPl'1l llll'aSlIl't'd l'oll!.thl\' b,' di"idillQ: I hl' total Yal'iatioll illtn It'll ('lass!;'}; ral1!,tilll,!- 'from t11(: \'('I'T 1ial'I'O\\'ly U-sha ppd ('olld i t ion ill (' lass 1 (Iig, ~a). r;)llIid most fl-(>(luPllt-ly in tlH' slIiJsIH'(,jps l'III'iill/-llilis, g!':l!lllall," opl'llillg up ill classes 11 t'o 1\r (li!.!:s, ~b-<'l. alld b(·(·olllin!.!: l'atlll'I' sho!'t. stOllt, alld l1Io(h'l'atl'h- lWllt ill ~'Ia:--st's YI io \TI ~ (Ii!!s, ~d-J) of IlIP 8011O/'{llsis rytlP. and j)l'()('l'('(lillg to a SIIOI'! (('lass L' \ '( Ii!!" ~1!') or IOllg (('lass X) (Iig, ~h) Bil'aight typl' ",llieh fl'l' ":tl'iation ill (\,[('b of nIP qll:lntitnti,'p ('hal':I('tl'l'!' has hl'l'n plo(t('(l in hisiogl'aills (Ii!!,s, :: to 7). \\'hi('h :11'(' _!!','ollppd for (';l('h slIiJsp('t'if's. sh()wing typi(':11 populafiolls and some uf till' IIlOI'P impol'tallt tl'Pllel;; ill nll'iatioll,

• THE XORTH AMEIUCAX SUBSPECIES OF CCLrCOIDES YARIIPE};-:NIS 7 • c

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1!'IGn(~; '.!..-Culicoi(/C8 ru/'iip('Jllli.y (·OlJlllll'X. dl'taili; of' palpus, 1III tt:lllla, Spl'rm:l fhl'ca, alld male genitali:l. a-h, HPl'l'lIHlthl'cae, ;;howillg" variatioll in ;;hnllP. :l("('ol'clillg' to ('}assl's, as lahl'lpd. i-k, ~[al(' g'l'lIitalia: i. ;;pillO:o;e 1Il'dNIg"ns of ,~onor('1l8i.~; j, gf'lIitnlia with 11I1I'ulII('r('s relllol'{'!l, ;;holl'ing- hnre nedeng'm;; and k, pnrlun{'I'l'S or 'F(II'iipellllil<, 1. ;\IUlldihl{', III-P, t'alpi: Ill, all.~/rali,,<; II, .wnorC'/l8is; 0, Oct'illf'll/aIiH; Ilnd p, '!:{lriil)('lIlIi8, q, Female :lntelllHl of (/'/I81/'ali,~, showing st:nsorin with extrfl s('nsorin pl'{'spnt on sl'g'll1l'lIi;.: I', I'r, alHl 1'1,1, • 8 'l'ECHi~ICAL BULLE'l'IN 1170, U. S. DEPT. OF AGRICULTURE

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O.LAM OW" STjL-LWAl't~ LBl

"U50UIIII pc T [IllS lUll ~ "LL ~"~ J" '0

.0

• ,4 LOUIS'""" ~'. n" "TON ROUG[ 1]' 1.IIIL.IO U -.1L1.0 II 1.4 I.' I.' 1.1 LL.10 II 14 11 o I • ~I J , 7 1\) .,. "lL SUtlORI" 1~' ••U"RCA AlOIA· ,UI AUSTRALIS FIGURE 4.-l!'requency distribution of six characters of Oulicoidcs val'iipennis allstralis. • THE NORTH Al\lEnICAN SUBSPIWIES OF C1TLICOlI)ES VARIIPKNNIS 9

'0-

COLOiJIaQO _ feln COLL,N! L '0 • II

,0'

.l L. 1:" t A t. I: T "11IItO II!

IT., ~ ~l ~ 10 SIN 1.0 U I," La 1.1 1.1 • 10 It '4 rutI 3 1 '" u U 1.0 '-' I""LPU, " 'JPf"...TttE CA IoEO'A 5U S ALBERTENSIS 1,'rntJIIJ,; :j,-Fl'f'ql.H'nCY dislribution of six clmnl(·tcrs of Olilicoi(/cs 'cudipenn is (/ I/ICrtCllS i.~,

JO- [J

10 1 r A l ~ ,.' CloLI,o".'A UU.,T ,It IfLIIIIt. IlTTIot[ -ri l1 L ! ~' U T AM All locallTI'1 n ~. l:dJ~ .. ACS a ~~ f['1", _UUVil.LI- iJI(U(Q A C1, I=ll=l' ~[5 ~A ... ~ ...... '

&111110 ....

g?!t:~·''''11110 ... 'Ol'" 10, IIIC_' CO .. au" caLICNT[

\.0 u: l4 LI II 1.1 • 10 II 14 17 ~[I '1 I 1 10 I I" "I 2~ U 3D S.' PAL'UI .," . SI'unUTHlca A[o(a ~-1L ;UI S Ol~ OR ENSI S

l!'IGUR~; B.-Frequency distribution oC six cbaracters of Olllicoilies 'variipenllis sOllorcllsis . • 10 rl~ECHNICATJ BULLl~T1N 1170, lJ, S, DEP'J:, OF AGRlCUIIrUflE

' .. LifO .... ' .. OftCUTT ~ d1 & LL ~" Jr 10· • "'T'"O COlUU'A.Cj • ..... LOO ... , { ~ A ~JJ 1=1 10· J

CAllrO'UII" HU[NllIIl A, 8" J1 lb [1Q, Jr jiI"

20·

CALIfORNIA eOltA. LAIC! 'OiL li!I2.22..Il.o 3, JLto 1.2 1.4 LII.I 2.1 • 101114 17 1 o r 4 j 1 5 T 10 S 'N PALII'US MUIQIIL[ SEN'OIIlA LJ 511'[1""THI[0& ..[Ol .... OCCIDENTALIS lUI

Ji'IG!lRg 7,-I!'reqnency distl'ibnf:ioll o( six charnetp!,s of UIlIif'oi(/cs '/,'([1'iillCllnis occidcllf(/li,~,

On our distribution mltp (Iig, 1) eneh poplliation of the v(tl'iipennis complex is definitely assigned to 11 subspecies on the basis of Itll characters shown ]n table 2, although in areas of intergradation a population may be so intermediate in ehal'actel' that this assignment may be Itrbitrary to It great extent:

• • •

""P:l l') o~ :::l

'""~ TABI,E 2.-Diaglwstic chaNlctC7'8 found in females of Bl108jJccics of tltc Culicoides Ynl"iipenniR comple,v, oased on ...,~ sel'ies f?'om, OJ' near, the type localities (oltm'acte1'8 of pr'ima1'Y i1np01'tatnCeitalicizcd) l') ~ 8 X~ (mandibb teet h) Xl (third palp!ll ::\2 (wing Icngth, X3 (anlcnnal ~ ~ensoria, Ilumber Meso segment, mm.) w length/width) additional) Sperma- Aedea- notal d Subspecies theca gus pat to w type tern ." Mealll D5 pcrcent ?I'lean /D5porccllt Mean/ \)5percent ::\[CHn -I 05 perC':!1l1; ~ limits limits limits limits Q ______1 I 1___ _ ?2 o II B variilJennis____ _ S. 06\2. 52-3. (10 . 1. 7(1 11. 53-1. f.)\) 0.11 I 0-0.110 ! 14· 44 12.11-15.771 J-lJ bare ".j australis__ 2.53 2.1\)-2.8711.5,1 1.3\)-1.{)1) 2.75, O. H3-4. 57*. 11..93 n.86-14.00 j II-llT. bare A (') sonorens£s ___ . 2.23 1. H\)-2. 4.7 I. 2(1 11.HH. 0U 0.831 0-2.01 1 ]2.17 \).04-14.40 I 11'-YIII I '~ln:noseC o 2. 2. 1. I S7J1:nosc .It t" albert.ensiL __ _ 40 13-2. 6711. 60 1. 30- 81 1. 78 0-4. 05'1 12. 22 I 10. !lCl-13. 48 II-Ill i H ocddentaNIL___ 2.2(j 1. 8!l-2. (13 1.63 1. 60-1. 6(; O. 1i5 0-1. 70 ! 14.00 I 12.10-15. no II-lII I barc C (1 I I o o t:l *Theor' 'ill ":lln(', nlthough 4.00 is thc aetnal uppcr lilllit pos;,;ihlp. w ;:

:::l,.....

~ t:l ~ 2:0- W

I--' 12 'l'ECHNICAL HULLETIX 11,0, (T. Ii. DEPT. OF MmICCI/1.'UlU~

Culicoides l'ariipennis variipennis (Coquillett) 1

Oerato/logon 'I:OriipC'/Il/i,~ ('oqnillNt, JUOl, l'. H. Sat. ~Iu". 1'1'0(', :!::: no:!; l'r:llt, 1007, 1:. ~. Dept. ~,"gr. Hill'. Ent. Bu!. (i-!: :!!i, Clllicoi/lcs eari.ipcuJl.iS Kit'!TpI', ]O()U, Gen. lll;;edol'lllu, fas(' . .J:!: :;:;: .\Iallodl, J,!J]:;, Ill. State Lab. ,:\'at. flist. Bul. 10: :!!J7; n(llTIIl:ln, Ill:!;), AlJlcr. Jour. Hn:. ;:;: 2S!); Whitehead, 1!l:~·1, Okln. Agr. gxpt. Sta. HIlt. 1!)a2-:H, p. 2(H; Hoot • and HOffman. 10:n, AllIer. ,TOllr. Hyg. 25: ];lS; l.'hoIII::en, 1 !)H7, X, Y. ( 'omell) AgI·. EXJlt. S'w. MeIll. 210: 70; Curtis, 1U.J], Brit. Colulllbin Ent. ~oC'. ProC'. :.l7: IS; Fox, J.!H:!, l'nerto Bieo .Tour. Pub. Health & Trop. :'.Ied. 17: .JJ.j; .Tohannsen, Ifl4:~, Ent. fiol'••\!Il!'r. Ann. ;~O; 7~0; .JaIlles. l!H:{, I:'flll.l':H'. 1';111. IS: HS; Vnrgas, l!l-lfl, Hc',,,lnsL fi:tlub. Ene. 'l'rop, ()It'xieo] (i: -I:;; 1{llIJ",JI'OIl and lJ'ronk, 1!)50, Kans, gnt, Soe..rOUl'. 23: 11:3; Knowlton llll(l .K:! rdos, lUi; I, Kans. Ent. Soc. :rOlll', 23: 103; Khala!', HJ(i:!, gilt. fiol', c\mer, .,"nn. -I:;: :{.J!J; Wil'th, 1fl5:!, ('alif. l'nil', Ellt:. Pub. !J: JSO, 2;)2; Ortiz and )lirsa, :t05:!, Al'I:1 Cient, VClIczolnlUl ;1: J:!7; !:-iIlOW alltl Pickard, In:;:>" Slll"'C>~' Artll. ClleslllPI' Creek, ~~VA HIlt" 11. ~I:;; I,'qote allll Pratt, 1!l;j.J, PUll. Hl'alth :'.IOIIOg'I', lK, p, ::,1 ; ];:l1l11U11(lS !llld. Keenl'l', ]n:;·I, ~rOslJuit() Xews 1.J: S::\; Fox, I!)[);), l'IIi1', L'Ul'/'tO Rico ,Tour. Agl', ;1!J: ~:;8; ('Oltl'l', Wil'th Hnd Knutsoll,.1!)[)i;, MO,;qllit" Xews]:;: 15;:;; ,TOiles, U)fi(j, ].;Ilt, Ho('. W:u;hillg'ton PI'O(', ;:iK: :!(j; 'Yil'lh and Boll illl('I', 1\)50,. :'.losquito Xl'w:; ](j: :![)()-2UU. Type8: To begill ",ith, we III11stdl'fine tht' population "'liidl ('oI'oad viUae. Scutellllm (fif!'. Sa) gray on ends, (-]1(' Iwoad, dark llH'(lian tll'l'H without II pttle spot-in t'he cent('!', 'Ving (fig. Ha) I>I'o\\'1Ii"h \\'ith dark markings dominant, the pall' areas slllall and dull gl'Hyish-\\'hitp: nw('mtriehia. mosily dark. ('\'('11 in tIl(' pall' ;ir('a~: ;l distinct nalTOW pale band OY('l' 1'-111 ('I'os;:;\'ein to mediol'ubital fork. (lis('al ar(l:t bl'O\\'I1' ish, the i wo djsi'al pa It' spots in (,(,1 I R r. ~eIXII':lINI. by a l)J'oa

1 ~ehe r(·f('r('l1!.'l's g-il'l'll herf' nppl~'l'o 111(' ('Ol/l)lll'x a:-<:t whol(', sC')lHrlltion in lIl1111Y enSt'S heiug' impossihlt' Of' hazardons withollt tlt(' ('x:lIl1inatiou of speCimens. • 'EHE NORTH AMEIUCAN SUBSPECIES OF CFLlCOIDES \'ARIIPENNIS 13 •

a varllpennlS

b australis

C sonorensis

d occidentalis

FWl"lt8 S,-('u!icoidcli r(//'ii[l(,lIlIi,~ ('ollllllc'x, wing and (l()r~al. tilor:l<.'i<: p:ltterll:': It, ·r(//'iiJl('IIIl.i,~ s. :4tr.: h. !l11~ll'lIli·' t 1I/1J!'l'tl'lIl!i~ liimil:lrl: 1\ XfillOl"I'/lM.~; d, o(·('itl('lItl//i.~ . • ].1 Tr':CHXlCAL B(~LL~TIX ll70, l', :-;, DEPT, !iF A(ilUCl'LTC·Rr.; g~J1italia,Oig, :2j-k) ,with ninth tt'l'gulll e\'pnly (,Olll' on Yentl'al sUI'faee of main body, P:1rameres with ~t()ut basal Hl'lllS, the ('n'e d.istal tips curw~\L latera]]y, then gnuluallr n:t.nowed to straight, slender: hyaline, simple points extending caudad, r-at'iation: The extent of normal \'a1'iation in the qua ntitati \'(~ ella 1' ;I('(('I'S of 'l'ariipcnni8 l'ariipcJlnis is ~hOWll in table :3, and the fl'l'qUl'IH'Y diagram:-> in Jigure 3, The wing length is quite \':ll'ialJle, the lllean of 1.0 111m, (){'('uI'l'ing in only :W Pt'l'l't'.llt of the s:ullpll's plotled ill till' histogl'tllllS; Ull' llUllllJ(>l' of nl:mdibulal' h'eth is only fairly diagnostic with thl' gl'enJpst ft'p(11'(' III (ill IH'r(,l'Ht of till' samples plotted; l\OWl'H'l.', olll,\' l~ Iwr ('pnl had wl'll-dpwlo)ll'll spitH'S, as in sonm'ensis, 111(li\'idnals :from thl' :-;outh('('n ~tatl'S \\'('re eonsistently smaller, with mean wing It'ugth I A IlIlI\. at ElIl'l'\'ilip lOkhhoma) , f,5 at Flat ,,'ood CAin b:ull:t l. alld 1.5 tI t }I.:\rianna (Florida), FOllr specimens from Rithland County (\,ris('on~in 1 with wing length of 1.·.1:-1.5 mm, wC're t11(' l'xeeplion ill the XOl'th. In tll(' South there :llso were -[ewe(' lllandihlll:ll' teeth. with a ll1l'an of' 1:3 ('aelL at Flatwoo{\ (Alnhnma). I [ll!:ilbul'g lTl'nn(,!:i~l'(,). allli ;'fal'ianll;l (Floridn). Loealiti('s with:1 ('ollsidl'l'flble IHlml1l'1' of extra sensoria pl'e::ent wel'e lhne~ Riehland, and W"nshbul'J\ C()uJlti~'s (Wis('onsin), Dubois (Illinois) l :md Hust: burg (TeJ1lH'SSl'l'), nil Oil tlIP Wl'stern edge of the l':lngC' of' the sub "peril'S, Habitat: In ,Vis('oll;;in tl](' junior autho]' reared /'Il1'iijJl IlIliN s, ;;(1'" from IlItHl and ('o\\' lIWlIUl'l' in a wet arl.'a. adjacent. tn :" waleI' tank in "rashl.lllL'll ('Ollllly, HIlll froJII tllp clay-loam 1l1:U'~in of Kn:l]lpS ('rel'k in Richland. COllllty, T\t(>I'l' is no ('\'idl'I\('(' that this sui>spC'l'ies is as~m'illt('d \\'ith ;->:dillP Plldl'Ollml'lIts. and this fnei.ol' HIlLy Sl'l.Te Pt\\'('('l1l'llJ'iiJi('llni,~ and its l'l'l:Ltin's is the scn.rc ity of' l'Ilriipul/Ils \rilhill ii~ rallgc, 'rIJis ~(':u'('itT ha:; hilltit'rpd Olll' study of sui>spl'cifh' \'llrintjoll due to the difliC'uH.y of obtaining nell' qllflte sel'i('s, )fa]]och (11:)]5) l'C('onh·d 1'l':u'ingv£Ll'iipcnni,'1 from :::ialt Fork at St. ,1'os£'1>h, Ill. Xot hadllg ('XHlllillPd this sl)('cillWll. we cannot lIi!:' ('ount the possibility of its beiJlg fluNtrulis, since a s:llinc enyil'OllnH'nt is indicated by the loe:dih' site, TIH'I'l,rol'e, this 'l'l'('ol'{l eHnnot be :lssignpd ddinftel.\' (0 a sllb~pe('il's, • rl'HE NORTH AMERICAN ::3G13SPECII::S OF lTLICOIDES \'ARUPI~NNIS 15

Distribution: United States: Altd'>flmll, Delaware. Florida, Georgill) Illinois, Inc1iflnn, Louisiana. :\Jarylnnd. ~Ii('hi~al1. ~[inn(,S()la, ~lis sissippi, :Missouri, Mont-awL, Nebraska, New .Tersey. New York, Ohio, Tennessee, Virgi nilL, 'Vnshington, ,Viseonsi n, :lnd ,Vyomi ng. Cnnada: • I~ritish ColumiJia. and Ontnrio, Earliest. recol'<1, IG ,Tnnuary, S:lY1UlIlah, 0:1.; Int('sl record 2Q Oc lober, Baltimore, :Md. Ten femrtles and 5 males taken at Marianna, ,Jackson County, Fla., ~[ny Hl55. represent a yariable population with ,,-ing measurements attaining extremes, two of the fn'c mnJC's }un'ing spinose aC' femaks nrlcl S males '('ollC'rtC'cl at KC'IT HesC'IToir. ~fC'cklenlll1rg County, Ya .. •\.llgllst 1()3·I,. 011C' JC'nmlc 111\S l)onl pnlpi with a nll·io of 2.5, and two maIl's han' about Gspines on the aC'dC'llguS. ThC' "'yoming specinH'n. ('on~i"til1g of thp male genitalia onl\". hft" tl1(' ae

;t is doubtfully l'dpl'I'(;d to /.tlJ'iip('}inis. H • Disl'II.'!sion: The suhspecies 1'{7riiJli'llilig i~ gPIlC'I';dly (li:>{rilmi:e(l O\'PI' the fClI'pstl'll l1orthen1 Hnd pasteI'll pol'iiol\!:i of )\01'111 Aml'l'ie:1. The wpstt'rll poplllnti01ls f['olll Y:L\'pnby (BI'itish COllllllhia). Pllllman (IYashington). ~t('\'('lIs\"iIJe tMontalla). alld H.o('k ~prillgs C\Yyol1l illg) in tIll' lI()rtlH'1'II Hockil';'; art' po:-::-:ilJly 1'('lil'{:, of n prp\'ioll:'. 11101'(' l'xtC'lIsin' 1':ll1gP dllring periods of gl'patpr IllOistul'P nll!l ('()OlH Il'm pPI·nhn·t'. IHJ:::-:ibly from Pleisto(,C'Ilt' linH';'. Culicoides variil)ennis australis 'Virth and Jones, new SUbSl)ecies

/Jitl[llI(Jsi.\': :-lizp mo\1:4 (rig. :!njl with third. st'.Q'1l1l'1l1 b!'()ad, :2.;) (:2. L-:Ul) ';3.1. limps :1:4 long as grC'n.tl':4t breadth, \\"ith n. broad, shallow~ partially • dh'i(lp

The type ,::erie,; from Hatoll HOll~!(' (Loui:-:iana 1 j:, 1'l'latin'ly ltonlo gt'neons. with slHll'p-praked frNluC'n('Y di,.,tTiIJlltioll:' for palpal ratio. wing length. mHndiblp tl'Nh. and antennal ;!rn;:ol'ia. Till' ;:pPI'mathe<'a • is quitl' nll·i:lhlp. fl'OI11 typP I to tnll' ' •. with a "light lW:lk at typl' 11. The :-itillwntpl' (Oklahoma) ;:('rip:, "hows gl'palPI' \'lIl'iability duc' ]l]'(' :;umably to intel'gmdarioll \\'ilh .~OliOl·I'I!Ni,~. a;: e\-i.\ I'allsas Xational ,rill/Ii fp Hdll!!l' \\,:l:-'; 1'(':11'('([ 1'1'0111 ,::alt pools ill n .....·,di/·o/·,da Illal'"h Oil till' tidal flat:' hOJ'(IL'l'ing ~all .\ 11 (onio Ba,'. /)i"tl'i/~/(ti()lI: ('lIit(,d :-;tatp;:: Kansa:-;. LOlli"inIlH. )Iis:'olll'i. Okla hOIl1H. ~ollfh Carolina. Tl'xas. and Yil'[[inia. Eal'lipst I'P(,OI'(1. 1 I ~fnl'('h. Pl'fl'l·sIHlI:g. ~[().: Intl':'t I·peol'd. :1 ])('('('111' UH, ('harlpl'tol1. S. C, )[tu'blp ('itO", Okl:! .. :'PPCiIllPIll' I'PPI'(''::('lltt'd by Ii f('I11:t!l':-; and .'-' Illall':; ('ol1(,(,(,p<1 lO~[n\·Ifl.):;. D. K Hom'll. illl'ludp (i lilah,s with hill'P apdl'a!!i nnd ~ wilh a fl'~\' spinps. . • THE KOHTH A~[EHLCAK SFBSPI':CIES OF C"('LLCOI DES '"ARf1PEKKIS :17

Of 28 femnles alld 18 males ('olleded at: ~tilhnl!e1', Okla., ill May alld .Tulll' J!)i);i. D. K lIowell, t'ht' femalps hn\"p a llIl'sollotal pattern between wl8tl'alis and sonm'('Il"~i8: and of the males. !) han,' a hare lIecleaglls and!) lmH' a spinose aNle:lgus. . • Disru88iol/. This suhspecies o('('upit's an ind(l[inite an'a ill the 10w('1' J[ississippi Vallt'Y alld (;ulf Coastal Plain when' /'ll/·iipnllJis. 801101'('1/. si,~. and ((lb('I,t('lIsi.~ ('onle to!-!l't 111.'1'.,\ ppa n'ntl.\' its partial e('ologil'a 1 is,olatioll ill salinl' ('n\'iI'OIlI1ll'nt's aids it ill nwilltaillilll! a distind popu lation ehnl'aetel'. Isolated SI)(>('illII'IlS 01' st'l'i('s \\'ith iWl'tial ('hal'ad('l' iSI ics of (l1/8tmlis O('C'UI' in populat'iolls of 80)WI'(-/18i8 in IIlallY T('xas l,nd Oklahollla lot'alitit's, slIg,!.!'('sting t hat the gellt'ti(' and. e('ologieal barrie!' to intt'rllI'e('dilll.!.' is I'platin'l" \\"('ak Iwt \\'(,('ll t'l1('Sl' two su\) spt't'i('s. Thel'l' is IIlu(,h' It'ss l,\'idpIH'(.: of poplllat ion illh:'I'nlillgling 01' interbl'('('ding betlY('('1l ((I/.I'//'((Ii-'< alld /,II/'ii/II'/1II i.'i, as al: _"al'ianna (FJOl'ida), but: 0111' sanlpl('s 1'1'0111 this pal't of t-1\f' ('ouiIlTY an' not su1' Ikient' to fell us llIu('h. ~[Ol'l' than any of' th(' oth('!' ;-;lIhs)lecil';-; of tIlt' /'((I'iipf'l/lIiN ('oll\pl('x, allstralis pl'()\)ahly shollld 1)(, ('Oll~,ndl'l'l'd a;-; all t'x:ullpll' of 'In '\'('ologieal 1':H'l'" 01' "l'('oty]>l'," altholl!-!:h g(l(lgl'aphic l'lellll'l1t;-; ilia,\' also hI' ('01\('('1'11('<1, p:->llt'('ially ill it':; l'l'latioll to 1/ Ibl' 1'/(' liS is. Culicoides variipennis albertensis \Virth and Jones, new subspecies

/)ill!IIUMis: ~iz~'I'l'lati\'l'ly laq!l'. wing Ui {L+-U)),':lT 1ll1ll, long. ('0101' l'atlwl' light: nH'sonol1l11l pal(" whitish gl'ay, pollinose with broad hilt: faillt \'pllowigh hl'O\\'n \'ittat': with 1\1I11\l'I'OUS, n'l'\, small d:lrk 1>1'0\\,11 Pllll(~tlll'l':;' of' \\'hi('h ollly a fp\\' in tht' IIlPdian :ll"('a tl'lHl t'o ('oall'sl't,. ~('lItel1ll1l\ blal'ki:;h Oil IIwdiall thil'd, sidt's !ll'(J:ldly y('l lowi:;h. Will/! ollt< (,«II:tlly pl'olllint'llt: 1l1:l('rotl'il'hin whitish ill tht' pall' aI'P:ls.lialtt'I' dal'k, whitish af 1>:1;;(' of sh?11l :uHl Oil 1l:lI.'l'OW hind Ill:l!'u:ill of kllob. _\ntt'llnnl spnsol'ia always jll'l'St'1lI' 011 ;-;('!-!:nwnts III, \'111-X (no t'xil'a S('IIl-'ol'ia, ~ !)(,I't'Pllt of' typl' sl'l'ips), alld sonll'tilllt'S abo 01\ sl'!-!:nl('nts 1\' \'11 (R pt'n't'Ilt). \'-I'll (I;) !JPI'('l'nt'), n-\'II (:ll 1)(,I'('('l\t). 01' n1 ollly (-P, pt'I'Ct'I1t), I':dpus with thil'd Sl'!-!:I1H'lIt iJl'oad. :2.+ (:LO-:2.0).:2'. timeg ag IOl\g as gl'('at('st bl'('adth, with a. bl'Oad. shallow, !:iensol'Y pit, nlOl'(' 01' lN~s ;-;lIbdi,-id('d into two s('pal'at(' pits. ~Iandibl(' with I~ (l()-I~~)/i)1 {p(lth. ~p(,1'll1at'I$'('a U-~h:lpt'd. IlIo~tl," t'.'·Pl' I. io tyPt' I\', wit'h a nl:ll'kl'd pl'o]lol'liOIl (:l:\ pel'('I'IlI) of I'ypc' IX stTaight, fOI'III:;.~lnll' gellitalia as ill /"I/'ii/)I·jll!is~. SiT., 1J1l! wilh ardeaglls ilt':ll'illg 1111111('1'011:; lill(' \'elllnd spi lies. 7'YPI'8: Holotyp(' !jl (pinlwd). Leth!n'idgt', ~\lhe1'ln, 22 .Tuly 10i);), .J. A, Dowll(,s (dt'positN1. in the' Canadian Xationnl Coll('('tiOll, Ot< lmyn). _\llof)'pl' 6 (slide), !:ialllt' llnt:l us tYIlt' (C.\X), Pamtypes, !jl !jl -;~;) slides, I:) pilllled: d d -:1 sli(les,;: pilllll'd: Ldhbl'idge, 10-2:2 .llIly I!mi), at lig-ht, .1 .•\. $o\\'nes. (n';X~f:!jl !jl -'I;) slides, ii pi1l1w(\: 6 d :2 sli(l('s, L pinnpd, l'l'lllaindel' to C.\X). 17 .Julle If););''i, !jl -I slidp (C.\X). Fol'l ~la('L('(J(l. 2:l .July 1!)i)i), swept • llwl'gin nlkalilH~ slough, .J. .\, Do",n('s, !jl!jl -~ slides, 1 pinned. 18 TECHNICAL BULLETIN 1170, U. S. DEPT. OF AGRICULTURE

J -1 pinned (CAN). Brook~, 18 .Tilly 10,'):3, at light,.r. A. Downes. 'i1 'i1 -1 slide. 1 pinned (CAN). Variation: The extent of normal variation in the characters of rtlbertensis is shown 1n table 3, and the frequency diagrams in figure 5. • The type series from Lethbridge (A.1berbl) is relatively homogeneous, with sharp-peakl'd frequency distributions for palpal ratio, mandible teeth, and antenna] se11s01'i:1. The wing length is quite \,:u'iablc, with a low peak at 1.u mm., and the spermatheca type SllOWS a scattered distribution. The Fort Collins (ColOl-aclo) series is quite similar to the type scries in all Chal'tlctetS except wing size, \yhich is slightly Iarf!t' I' (lllt'an l.A 111111.) and the malt' :lPdeaf!us in w11ich three of seven Bpecimens exami1H~d showi'd fi'w or no yentral spines, indicating probabJe intergraclatioll with /'(ll'iipr'nnis s. 8tl'. Other series are too short to indicate distriblltion oi' characters. Habitat: For oUl'information on the biology of albe1'tensi8 '\'e arc indebted to ,T, Anthony Downes (in 7itt.) who \'ollected t11e type series in Alberta. They were taken at the ll1iH'f,!:ins of the allmline cattle pools, whkh are a cluu'acteristici'('atul'e of the nortlwrll Great Plains. DistJ'iblltion: United Stnf"es: Colorado. Kam;as. )[ont.aIltt, Nebmska. Oklahoma, and South Dako(-a. Canada :;\Ib('l'til. . . Enl'liest reeol'd, 2G ~\pl'i1, Lineoln, Nebr.: hie::;/: ]'Pc.ord, 28 October, Fort Lupton, Colo. Nine females colleci:('d at Fort Lupton, ,Yeld County, Co\o., 28 ()ctober 1!)40, from :l window of a dairy b:ll'1\, by C. 13. Philip, are vcry large and dark, with well-marked brown "ittae OIl t11e lllesonotUJl1 l'i'sembling the Utah populatioll of 80nOI'N!8i8. Di.~(,lt8sion: The population of Cl1l8li'a7i8 found on the salt marsh at Aransas National "Tildlife Ui'Juge on the Texas Coast <1iil'el's wry Iittle from albe?'tl'J/8i8 i'xc('pt in the higher proportion of specimen's with bare aedeagus and the presence of more exh'll sensori:l on thl.' i\'mall' .lllti'1I1wr. \\,itb sueh lillking popuhtiolls still pxt:ant. it is reasonable to postulate that tl salt-tolemnt, (lu8{I'alis-s01lOl'r'IlBis-t'ype ancestol' may han~ become fa irly we1l eli f1'prpntiatpd in the North. and al::;o nUl\' h:l\'p !!L\'PI1 l·i:;(, to Itlb('l'!('II.~;s in thr alkaline sloughs Rnd pools chamct·l'l'isti(' of thellOl'thi'I'!l high prnil'ips at the ensh::rJ1 haf'e of the Rocky )[ountains. Tn the South there has been no isolation from ausf1'alis and 801101'1'I1Si8 and tilerC'. is at prpsent an area in the southern Great Plnins wh(,l'e a pool of Yal'iability stilt exists, and sppciJl1el1s cannot drlinitel,r l)(' a::;sigl\rc1 10 slIhSI)('cips. ,rhrlhpr t1w intl'rgl.'fldntion will pl.'O"e 1·0 he geof!l'aphically and genetically dinal, 01' \\'hr(llel' morp compl('x prtf:el'l1s ('xist tkpellclent 111)011 IOCf1.lpcologi cnl eonditiollS, can he detl'l'mined onl\- hy flltUl'(\ intl'IlSi,'p collection nnd biological st.udy. . • Culicoides variipennis sonorensis Wil'th and Jones new subspecies TJiag·nmis; Size sma.1l, wing lA: (1.2-1.G)!17 mm. long. Color .light; mesolloiu111 (fig. Sc) pale prnillosP. gt'ny with llOt VC1'Y numerous, large, blackish punctures which uSllally coalNlec in f,.roups of two to flvi' and for))) lal'gPl' spot's, ,rrrqllrnt·h· characterized by three distinctive tnms\rerse grOllj)S 01' coalesced SP~)ts, usuillIy no tl'nce oi~ • brownish yjttne on the gl'ayislt background. F\C'lltellul11 ypllowish 011 'l'I-m NORTH AMERICA);' SL'BSPl·:crES OF CGL[COIDES \'ARllPENN1S I H

sides, blackish 011 medinl thinl, lI~u:tlly vale Oil hind IlI:U'gill in till' center of the mcditlll dal'k aJ'l':l, Posts('uteUlIlll dark gray in middle, whitish pollinose on sitlt'S, IYillg with light and dark markings aoout equa.lly prominent alld nppl':ll'ing delinite (lig, l)e) l dark spot on base of "ein ~[2 shol't to long: nlll('I'otriel\itt whitish in the pall' al'en~. • Halter whitish. onl r the base of the knob dal'k. Antennal sellsol'i~l ahnl,Ys l)l'l'sent on st'glllL'nts JlI" rIll-X, and. sonll' times ahio on seglllPnis \']-.\',[ (-1:-1· prl'cellt of tYPl' seI'les) 01' ollly 011 Yl.I (5xi pereent). Palplls (fig. ~n) with third ~egn1('nt, broad, 2,1 t L!)~2,5)i 12 times as IOllg as greatest bl'eadth, WIth :t htl'ge~ ~llttllow, inl'OIU plctelv eli \'ide{\ sem;()I'Y pi t. Manti ible with 12 t 11-13). ID teeth, Spermaf.hee'lL shol't :lId stOllt, :tln'upll,)' bl'nt in ll1iddll' (t,rIlt' IV 10 \'1). Male gl'nitalia as in sllilspP('il's /'w·iijiUtJli.l', 11I11 wilh aplopl'd \-t' II I ral "piIH's OIl maill body. Types: Holo[ypr ,; (:::Ii("[('.), SL David. (\wlti;;p Coulliy, ,\riz.. Odobl'l' lHii:3, light irap. r. S. PuiJlic .I [l'tllth S(,ITi('(' (t.nll' :\0. U:l2-W, l-. S, X . .\1.), .Alloi.\'lll' ¥ (,.:Iidl'). l'aral,nle,,: ~alHe daia a;; t.nll', datl'S Sl'ptpnlilt'r· Ol'ioiJPI' 1\1;1:3: if 9· t:l ..;Iid(>s; ;)1 pilllll'd: 1 d -IU Rlides, ruriation: 1'11(' norma I. \'U ria j.ioll ill t Itt' ('haruet('r,; () f .l'OllO/'CIt8i8 i: showII in table ;\, and [he fn'quPl1ty diagruIlls in figure G. The nlria [ion is l'X(Pllsjve and diflieult io :1l1:1lyz(' bC(':l.llSl' of shortage of mall' rial frolll St. T)avid, .\riz.. ihp IYPl' lorality. ][O\\'CH'I': the serie: frorn. fin Arizona. lOl'Hlitip::: (\\"('\Ilon, st. Da\-id, .I [ot .springs, Oak Creek CaJlyon, a.nd Sabino ('all,Yon) and from Agua C:diente ,Sonora. ~[exieo), appeal' to be I'll tlll'r h0I110gl'IH'ollS alid Ila \'P. bl'pn ('0III bi ned for IHlrpose.s of unal,)';:;is of lloL'lllal \'ariatioll. .In I hese combined seri(':S [her(' art' 8hnrp-lll'akp(lfn'qul'Ilt',\' distrihulions for palpal ratio (mcan ~.21, wing length (1.:1 nlIlI.), mand i bl('. tppi It (1 ~ l, nlld l'xtra :tllil'llll:t1 sPllsol'ia ((I.D)' wit It lL 10\\', Iwoad ('Ill'\,(' on Iy for :-;p('I'nliltlwC:I tYlll' (Ill· \~I1). Hut ill dilrVl'Pllt portions of tllP rallgl' S(}IIOI'{,Il,~i.~. \'ari ons eh:traett'l's ;;hift lllarlwdly In Jrequl'uey. l'lah SpeCillll'nf> al'p larger (wi ng l.(i-~.tl 111m. long) wi tll 11l0l'C lll:tnd ibll' teet It (1;3), pos sibly jllllieafing intel'gradation with l'Cll'iip"nnis s, stT. ",Vnshingf"()n speeinH'ns tt\'('. indislinguishn.hle from (}(·(·idl'/l.faliii (IXCl'pt by till' ('hal' ndpr of the spillPs on tIll' mall' aedpa)!lls. ..\Iany 1\'xns. Oklahollla. and Ka1l8:ts sjl('eilllPlls show signilieantiy lllore exil'a sl'llsol'itl {mean lA). thus indieating inll'l'gl'adatioll wit·h a1l8tl'ali.,' 01' albel'tnrsis . .A long, typical, 1IOlllO)!l'IlPOU:-; sl'I'il's from Big Bl'IHI Xnt iOllal Park. Tpxus, was llsP(lfol' the slatistical llllalysi8. The l'platirt'ly great. nlriation of l'hnnH'iPI's ill till> Texas Sl'I'il'S or S(}]WJ'Cl1,~i8 lllay IHl\'P SOI1I(> :'(,:IS01l:t I eOITelntion, "rini('\.' fOl'llls of SO 11 0 l'('IlHis lIPI)e[tl' to be t1[lpl'pciahly ]:u'g('r Ihall thosp ('olll'dedin thc' \\,:11'11\(,1' months. SOl1l~' spPl'inll'IlS t:nkenin Mal'ch at Kerrville definit"ely l'esem ble 'l!al'iipl'nnis in color ma.rkings and struchn'ltl ('htl r aci('rs. KeJ'ITil\e light Il'npstook It considerahll' (lI'opoI.tion of speci nlen!) l'l'se.mbling m{81f'll1i.~ in Mny and .Tune of H):H, but not in otitl'l.' monih!). Otl1l'r ;,;pC'l'inll'ns similar to (l.!lstl'alis wem taken with 801lOl' l'I1Ris at. Canlp Slnnl!',\' (Octobpl'), Dryden (May), and S:H\(It~I'S()n (April) in Tpx;18, and at Enid (Julle), Oklahom:l City (Allgl\sfL IIIHI Hf"illwatl'l' (May, June) in Oklahoma . This apparl'lIt illtergl'tltlalioll of sonm'cn.si8 with 'i'w'iipcnnis a.nd • aw3{m/i8 is prohably du(' 10 1'!'l'O\llhillaiiolls of chal'af'fC'I'S of Ihest' thret' 20 TECHNICAL Bt:LLETIX 1170. r. S. DEPT. OF AGRrCeUl'UBE subspecies in a heterogeneous gene pool in t-ll(' zone of oyerlnp. Thl' fact thntin difl'erent, samples there may bl' difl'l'I'l'llt ('omhinat:ion~ of characters showing illt('rgmdation rule:> out an)' possible int:l'l' pretatioll that ~'elleti('all'y separacl' POPlIlHtions ill.·e in\'(}ln'll. • 11abitat: In Texas. NOno)'('/lsi8 was Illost' t'Olllmon h· 1'ollnd in low to moderate, density in . lllUd. at the Illal'gills of stot'l( ponds and slow streams where there wt\» nlon' 01' It·s:> pollution from sheep and CO\\· manUl'e. lIowe\'(~r, we haw 1'olln<1 8()1I()I'ensis hl'pe(linp: in greatest density in pollul'!'tl mild in se\\':lge (lflllIl'nL ",\I' Paris (TennNi»l'e) Snow l'e:II'e(L~onOl'ell.~i8 fl'Ol1l a pastu l'l' ~el'p, ~\t :--ihafh'r (Cal ifornin) Brookman real'ed this »ubspeeil's fl'OIll a IIl.llnieipal 8('W('r fal'lll \\'I)('re sewagn 'watl'l' \\'as uSPd for il'l'igat'ion of C'rop8, 1Yhih,lwad (l!);H) I'pporil'(/ a t'l'lll:lrkahll' outhreak of /'((l'iip"lIl1.iN Ilpar th(· XOI'lh ('anadian Hin'r at :--iha\\'IH'P. Okla. 1"0111' slidp mOllntpd malesof1Yhitelll'ad'» lllatpriai in thp r,:--i. Xational )l'IISCIlI1\ sho\\' two with ban' aerleaglls (/ /l/lsh'aliN) and 1\\'0 with H'ntl'al spinet' (albel't(,/I.~i8 01.' 8(}11OI'('IISis). 1Vh itt'head t ra('C'd I'll(' SOU1'('l' of the in fl'station to till' riwr. whkh at that' tillll' was 11 sluggish stl'l'a11l a,1'l'w ranIs wide with a \'('IT sma II \'0111111e of wal'Pl'. The \\'atpl' was hp:l\'ih' j)ollllted hy Sl'\\,agl·. ~\'h ieh was d llnl]H'd into the ri WI' at Okla h0l11~1 City, on'l' i'OI'ty 11lilps abo\'(' Ow olltbreak arpa. 'fhe scwage tlesh'oyed pJ'aC'tieally all allintal life in nil' sll'eam for many miles dO\\'I1»tTl'1Ull. "U Shawnec tile stl'PlIm 'had rp{'o\'('I'ptl enough to support a trell1l'l1dollS ('ulit'uid('8 po])ulaJion bllt yery litlp other animal life. Adult ('u/i('oides \\'el'e partiC'lll:lI'ly anlloying to li\'cstoek but did 110t attack humans to nearly so ~Tl'at an I'xt'('nt: :,IS onl' WOllld C'xpcet f!'Om their numbers, l)i.~tributi()lI: l~nitl'd States: _\rizona, ('alifornia, Xevada, Npw )(exieo, Oklahoma, Telllll'SSpe, Texas, rtah, and 'Yashington, ~[exi('o: Guerrl'ro. )[exi('o. D. F .. XUl'\'o 1.-('011. Pllehla. nnd SonOl'a. Earliest re('ord, '1:2 Febntan'. ~\ilstin. Tex.; .1af:t·»i I'('('oj·d. XO\'emhl'r, Patauonia. ,Ari~. " " One of two lllall's ('ollp('ted. at Hakpl'»field, ('alit, ~(i .Tilly H)+G, H. E. :McClure, has it hare aedeagus, aml perhaps i» o('('identalis, as also 1 of 4: males ('olleeted at :--iangcr, Frp»llo, County Calif., T. Haley. Female speeimel1s ('ollee/'(·(l on \'arious dates bptweell J9J:i :lncll!)+S at Indio, Hi\'erside COHnty, ('alif., nppearindist:illglli~htlble t;-onl Imperial Yalley occidenta/is and may l'('pl'esent all inft'l'gl'Hdation. FOlll' IllU les co\ll'dl'd at Oklahontn. Citr, Okla., ~\llgllst H)B+, by F, K ",Yhiteheatl, HI'P, IWl'haps intermediufl, ((1I8tl'alis, :2 l11aks ha\'c bal'l' aedeagi: ~,spinose. Charactl'I's of ~ malps with bal'p aedl'agi and J female il'OI11 Paris, Tenn., .luly 1\);'j5, l'eal'ed hOlll pa~tul'l'; seep by 'V, E. Snow, »how intel'gradation with l'C1I'Upellnis. The following" variation was fOllnd in »I)('t'illlens from Texll», .~ femall'S Hila 1 male. of it series oi' a.J, females :I,nd 'j males frolll Calllp Stanley, Bexal' ('ounty, 2Z Octobel' ]0:>+, I'csembll' (ru,~tl'ali8; olle 01'+ female'S C'olleetecl at Dryden, 12 )[ay 19M. O. Sc'hollliJerg, resembles (lIl,~tl'ali8; 8 fl'mall'~, of it series of HH fl'llla Ips an(l 1n 11lal{'s fl'om Kpl'ITi Ill', U>ii~\ an(l 1nri+, T.J.•r. Bottimer. rl'sPlllble (l1l8ti'a7i.~,' 2' male;,; and 1 fl'malp. of It scrips of to females and G males l'earp(J fl'om sewer l'mllenl', Kel'!'\:ilh'. :~o )larC'h 1955,11", ,Yo 'Yirth, resl'll1hl{' I'ft)'iipt'llllis: and 1 of 4: fl'males culll'('iecl at Sanderson, 'l'l'1'1'('11 ('ont,\'. ~s .\pril 10;,)+. O. f-\('h0111berg. l'Psl'll1bll'R (1/1.~t/'{l7is on nlPsol1ohll paHprl1. • Culicoides variipennis occidentalis Wirth and Jones. new subspecies

IJiagno8i8: ~iZl' llIodrratrl.r1:11'gr, wi ng- 1.G t L()-,.l.1) i;~:J In Ill. Iong-. • Color light; meSOJ\otlll11 (fig. Kd) pah~ prllil10sP g-ray with numerolls small c1a d~ brown IHIllClli rrs tlia I: 'hC'<[uently ("oalr::'e(' in grou ps of two to lin', TOlTlling- Jarg('r i:ipots, r5pr(:inl1y in tile medial area: tilrer moderately promin('llt, pall' \)1"O\\"nis11 \'ittnc wmnlly pl"es('n! Oil tile fTrayish baekgronnd. :':klllelllllT\ \'pllo,,"ish 011 side,,;. bln("kisil ollllledial thil:'d. with l'l 51llall wedinll pall; ,..:pot on hind l11lu'gin ill tlie center 01' the I1wdian dark area. PostSt"lttplllllll claTk gmy in Illiddle, whitish Oil side's. "'in!:r (fig.l-)(l) with light all(l dark 11l,lrkillgs sharpl\' eOIl t raslillt!". til!' \l'II'it isl\ nrea:; 11101"(' (:xtl'llsiw titan tll(l ([ai'l: OIlt'S, :l1'ea, in ('I'll H;' IX'hilld tit(' 8('('011(1 mdial ('ell 5I'areel.\" dltrk~'I1Nl; dark spot Oil IJn:ip or \'(·il1 Mz Wl'y shott: 1ll:ll'I"otl'iehia whitish in tlIP, p:tl(· areas. llaltcr whitish. base of knob (lark. _\llt(,IlI1:11 S('IlS(lI'ilt ;dways pn':'('II( Oil :-:~'~IlH'llts III, nu-x (no l'xl I'lL sPIl:-:Ol'i:l 011 ij(j pl'I'e('nt of f.,rpl' SPI'jI'S), aml sometime:; :t1:-:o 011 \ VU (~ IWITPIlt), \'1-nl (t; !lPJ'('Pllt) 01' \'11 ollly (;~(j pel't'PJlt). PalplIs (fig. ~(l) ,yith third spgll1(>lli t)l'on.d, :2.:3 (1.\)-:2.3 )/:2~ times as IOl1g as !!"I'PHtpSt bl'l'adth. \\,illl a I:tl'!!"l'. d~'('p. H'll:;OI'y pit lI:,wtlly (1)('11 illt!" by a smaIlPI' \)(ll'(" ~'rn.ndibll' "'itll H (lJ-Hj ):ri tl'pth. :-:lper- 1I1;~tb(;I'a lIswtlly .OI1g- al1d broadly U-,,;]w/ll'd. l,r]l(' III'")...\lnll' g'er)italin as in :-'lIb:,])('('ies 1·(II'iijll'/llli,~. the HP([pap:lIs ",il hout ventral spJnps. T1IIiI'~: IfolotY]lt' ,~ (slidp). BOl'ax Lukp. Lukp ('Ollllty, (':tliL :2!> .Tuly I().j.s. 1'('al"PllfI'OIll lakp nH\l'gill. "r. '\". 'Wirth (type So. G3:230, U, ~. X. )f.). ~\ll()typp ~ (pimwd). :-:anl(' data. PnratY]lPs, samp data: ;;, :27 piIlJH'(l. 6 0 --15 slides, 1·1, pil1ned, ill alcohol-many .3 l's of (J('cidf'lI/ali:s is s]lOwn in tahle :3 find, the 'fl'pC[lIPJley sf'I'n :;(,II:;ol'in J. TIll' spel'mHtllPc:t is (juite YHl'inlJll' fl'OIll tYlll' 11 to IX. with a ]ow pl'ak at type IY. The s(,I'i('s ['!'O1ll Snl'nloga ~pl'inp:s ('alifol'nia.) Oil tlw pastc1'Jl boundary of the range dilrcl's mainly 111 Iwing of smaller sizp (\\'ing 1.3 mill. long), aJld. h(wiJlg 1'l'\\,el' mandibu lar {(,pill and mOJ"l' :;('nf'ol'ia~ indienting- ,])o:;::;il>](' intel'gradn.tion with 8onoi·eI18i.~. {Belkin Hnd McDonald, 1$)5(;, ha\'e described tltepeculiul' i'eatul'£'s of (hi::; locality.) The Kall1100ps series from tlte llortltel'll end of nle I'llngl'is YPl'y ;;imiliu' structurally fo tIle typl' series from Borax Lake, but !'Ite black pigment oi; the wing and thol'neil' markings is much mOl'e in«'l1s(', possibly due to an el1vil'onment:ll influence, ' Till:' scri('s fI.'Olll HII('IIrnW and Y('ntllJ"t\. in th(' CllJi:fol'lli:~ m:tl'itill1r mnl', shows mal'k('d (\itl'erl'ncps. slIch nslnrg-('l' 5iz(> (\\'ing 1.S-2.1 mill. long), more ('sil'n sl'nsol'in. (lnd gl'I1(,I':llly d:u'kl'1' and mOJ"e ditflls(' body marking-so )<'\ll"Illel' collecting max ]lI'O\'(' Ihnt this population is ('111 m:tl'cl'ia.l ([OP:-; nol' justify sll('h n. liJ1(', divisioll • al' thi$ time, 22 TECHNlCAL Bt:LLETIX 1170, L :-;, J)I':PT, OF A(;HI('n/l'lTBE

.\. compal'iSOll of the dat(',.; of ('alif'ol'llia ('oll(,<'tions SIIOW" no eol' relation between season (ti 111(' of ~'E'a 1') a nd body siz(', Ruth Hi' was reflected in the 8onol'e1!,~i8 population in Tc'xas, Xeitl\('1' WerE' t11el'(' any appa1'ent seasonal di ff(>l'el1eesi n th(' f'I'(>C(II('IH',\' of colledion be- • tween occidentali.<; and 8o'1U:n'('71sis in California wh(,I'(> the two suh species occur in the same geogl'apl1ie arpa, [[abitat: TJle t'ype ~'I'ies was I'('a 1'(>d fl'omt hl' 111 a 1"I.6n of a large nlkaline pond nt Borax Lakl'. Calif. ',,'il'th also I'l'al'l'd o('eidell/a/is fl'OIll puddll's polllltr(/ by lin'stoek ('x(,I'pta in till' Illal'gin of' a small ('I'(>ek at Ol'elltt. ('alif, Bl'ooklllilll 1'e,II'('(1 lal'~(' IIUlIllll'rs of llt.is sub sp(>cies from the mal'gins of 1'0;';0 ('I'(,pk, ('HI'I'.\:ing salt \\'atl'l' frolll the oil fi(>lds in 1\:£>1'.11 ('olmtT, ('aliI'.. and frolll pollutNl walel' at the hewageil'l'igation farm at' Hhaftl'l' in thp Salll(' ('ounty: ,\\'i1'th rpal'pd tllP 1:1rge da1'k JOI'Ill at HIH'npllIp. Calif.. in til(' ::andy margin of a s:t1t mar~1t pond just ba('k of the Paei!i(' O('('all I)('a('h. I n all tht':il' habitats the lal'\'tll' and pupae O(,('IIITNI in tl·(IIll('IHlolI:-:llIlllllwl'l-. :forlll ing almost a solid jine 01' bnnd a( till' nlargin of thl' watl'r alHl tnlld (ll' &Illd. Til thp Colulllbia Hin'l' Basin in ('l'ntra1 "~nshington. Bacon (In;:;;~) (ook orrir/('nlaiisin lal'gp IlU!llilHS in the. saline Illtll'shp!4 all(1 ponds at' lhe Potholl's 11('al' <),Hulli\'tln Dalll. Hpeclnlpns takPn at" Hoek." Ford ('I'pel\: and Lowpl' ('mil ('I'l'ek in Ih(' sanle watel'shed \\'PI'l' 801l01'(,118i8• .\('cording to Ba('ol1 . .:lnoplie/f's f,'er/Jol'ni _\.it'kc'n w('r[' takpni'I'ol1l thp lattpr two ]oealiti('s but· not' from th(' Potholl's. and Wl' lwlip\,(' tile "ame p('ologica} separation. flJ'olJabl,\' a l1laUpr of snlinity. n('('ounts 1'01.' the g-eogrnphieal separation ]1 e1'(' of o('(:-irlel1t((li.~ and M)]Wi'eIlNi8. Bac'on giws the i:ol1owing- notes on il. I'pnring, probably I'pfet'l'ing to o('ridentalis: ·'.H saIinc Scootenp\' La k(' ill Fl'ankli n Count\' tlH' waipl':-l \\'P1'(' Jound. ill .Juh'Will. to ('oiltnin ]wll'id lan'al', which s(>('Ill(>(l to share zootic dorllin'ntion (If theil' plwironmpllt with the lan':1e oJ an ephyc1tid (shol'eft."), .Adults of ('lIli('oir/r8 l'((J'iil}('ll1li8 (,Illprgt'd 1'1'0111 samples 01: lake Wail'!.' l'ptninpdin I'('al'ing jars." /)i.«tributioJl: {'nitl'd Htatt's: _\ rizolla, ('nJiforli ia, Or(,![Oll, ~lnd ,,\Yashingtoll; ('mlada: British ('ollllllbin: J[pxi('o: Bnjn. (,;t1if'orni~L. Enrliest ]'e('ol'(l, F('OI'IIHI·Y. FOl,t 'Yuma, Cali L: lalL';..:t· I'('cord, (i Sl'P tembel', Knrnloop~, British Columbia. Two mall'S of a, seri('s oJ U mal('s from Blythe, Hi\'('rsidl' ('Ollllt\'. Cal if" posse'ss spillosl' He(ipngi rescmbl illg SOIiol'l'lisis. _\. ~'I'il's of ~l females and 1~ male::; Jl'om l[uelH'llIp. ,\"pntllra ('ount\'. ('alif.. 18. :W ,J unt'. ID,IS, 1'(':11'('(1 frOtn a sa IL nHtl'sh pool. '\r. \\T.\\'i I'tli:, a rp \'PI'Y l:;rge and dark. In a.sprips of ;llll:lles frOIll Hpst'ing- Spl'ing~. 111,\'0 Connty, ('aliL, 20 )Iay 1%;,), .1, X. B('lkin, 1 malt' with spinose :wdpagllsl'C' sl'mbh's 801l()1'(,iI_~i8. One :tpma Ip 'from Sania. Cl'UZ, enJ if.. .Tilly 19+8, JL Coleman, is In.rge and d(ll'k as in the Hupneme spries. Two f'emales and 2 males :from Ventura. Calif.. ]0 April 10+S, H. Coleman, arc> vely large an(1 clark colored Hpady as inv(fl'iipellllis and are doubt fully refel'l'C'd to occidenta/is on fhe hasis of (listl'iiJut ion. • • •

'.I'A HI,}; :).~·Jfr(/811r('m(')1l8(~f/hf 8!1b.~]Jf'('i(,8(!f I/;f Culicoidps '-HriipPllnis ('umple-./! (Wi1U'S ]I"(,8(11/(({ as "m('an (m.inim111n ~ 1I1(l.rinl1lm) J/111m ()f'" __ ,,,, -+W.p.-._ ~

IAIIOIh! willtIt .11ill ililctt'rs S1tIllIJ/'r.~XUlllbcr Cllllada, Brit.ish Cohllllhia: i-:\\'Pllhy 2.IH2.S·:).1 );:3 2.0( I.S--2.1 J/·I 1·1( 1:1,15) i 7 OJ·1 ") ;~.. '1\'i~consin; y, Dnne County [2J I, ... :t:H:t I-:U) I:) 1.8( 1.·1·2.0)1,1 1.j(12··!fiJ/5 0.2(0-1)/·1 :r. Grnnt County 11]. :t2(:t 1-:t·I)/2 1. 7 ( l.i') /2 15/1 014 j Richln.nd Counh' _" . :t I (2..1-:Ul!-l l..If 1.4-1.5)/·1 15 (J 2-17) ji' 0.5(0.2)/8 ~ Rusk COUII!.Y [5L.. __I :l.2(:l.()-;).51/{i I. 7( 1.5-I.Sl/O 1·1(12-15)/10 o/n ":i Washburn Coullty [a] · . :t2(2.R-:l.7)/4 I.S( I.S-I.D)/:1 \(\( 14- 17) IG O.:l(O-I)/n Illinois: Dubois [71-.__ ... .[ 2.!)(2.li·:t2)/O I.U(1.7-2.2)!I() 14( 1:)-15l!B 0..1 (0-2}/lO ~ __ ::: 2\\issouri: Kirkwood , ...... i :3.0/1 2.1/1 15/1 , 0/2 :r: Ohio: Lockwood ______...... _. I I a.:3f1 1.0/l J.I!2 0/2 o :-rcw.Jerse\': Ncw Brullswick .... _. · .[ :tOjI 2,2/1 14/2 0/2 Dclawnrc:' Donn Lanriing.. _. _ . · ._ a.o/1 2.0/1 15/l 0/1 (') :\larvlnnc1 : '1 1~cltsvillc12]_ .. _____ • __ .• _ '. I 2.S(2.7-:3.0)/:) I.S( I. 7.. 2.(l) /:3 1·1( 1·1-15)/4 0/:3 Brinklow [4]- ______...... _I a.2(:3.0-:t5l/11 1.7(UH.S1/II 15(1:3-1(;)/7 0/1 L ,Tirginia: 3 Mount Solon [5] ...... :1.0 (2. S-:l. Ii) /n I.7( I.7-1.8)/;'j 15(l·HOl/O 0/7 '-' ParnRssas [31_. __ . __ ...... 2.S{2.7-3.0) /a I.7( I.7)/:3 J5(14-ltil/:3 0/5 ~ Kerr Rcsvr. [41-- ____ ._· ••. _..... _... ;3.()(2.5-3.3)/S 1.7(UH.S)/·1 J5{14-1Gl!G O/S ;' Georgia: ;.; i\Illcon __ . ______. ___ .. ,_ .•.... ;:::) :~.O(2.S-:3.:l)/:{ 2.1 (2.0-2.1)/3 15(1:3-15)/4 0/.1. Sllvllnnllli______.. .. I :1. I (2.S-a.5)/G IA(1.2-1.5)/i4 14/1 O/G "C t=i Alabama: Flatwuo(L._ ...... _I :l.l(2. ;)-:3.~)j(i 1.5{1..1-Ui)/G 1:3(11-15)/10 O/S Tcuncssec: Hustburg ... __ ...... a.2(2.~-~.0)/IOI.G(1.2-1.!)/la I:HII-l5}/ID 0.5(0-21/21 Z I Z Ji'lorida: Mariannll _____ ...... O' _ 2. S(2.0-.1.1) /·1 1.($ 1.5-2.0)/S 18C1()-15)!1:3 0.S(0-:3) IS ..... __ __ ;:n Louisiana: KilbourTIP ..• .... ~.$(2. ~-:l.7 ) /(~ U\( 1.5-1.0)/10 14( 11-15)/12 0/11 Oklahoma: El1crvillc__ . _... __ .. _.... _. __ . I .3.0(2.1-.3.2l/h 1.4(I.:H.01/7 1·1(12-15)18 0/0 ~ I Kllmbcrs ill brackpt,; rl'f!'r to Illllll\wr of Hpecillwll:; indlldl'd in st.atistical :lIlalysis, SI'C p. 28. w TABl,);] 3.-]t[easu.remenfs (d the subS1Jecies of thc Oulieoid('s vnriipl'nnis cOml)ie:); (vobles pl'csented as "mean ('Ii~';:nimllmI.\:) '/il(/,j,:im'll,m.) /nu'mbel' oj mea,s7ll'ements")-Oon ti n ued ;j:;.. CITI,[(,(llDER VA1UIPJ;XXIS Al:S'l'IlALIS ...:; M o Third palpal Teeth on Antennal ~ Locality !7. segment; Wing length mandible scnsoria .... o additional ;> t" tc Length/width :lFili1:meiers N1I1I!ber.~Number ~ LouisilUla: ~atonRouge [281___ --_- -- - ____ -- -. -. ______.. _I 2.5(2.1-2.9)/35 1.5(1.8-J,9)/3!l 11 (8-14)/Gj 2.5(1-4)/71 t" South Cllro!tna: Charleston __ ,.______. ______.. ,. ____ ' _ 2.1/1 ______.. ______j 1(11-12)/2 .. '. 2.5(2-3)/2 t: Virginia: Saltville ______. - ______._. 2.11(2.'1-3.2)/15 1.7(LG-L8)/7 13(11-15)/9 "-J _: 2.1(0-4)/15 Missouri: Pctersburg______. _____ 2.G(2.3-3.0)/24 2.1(1.11-2.1)/12 12(10-15)/22 ...:: --- 3.5(2-4}/23 '"' Kansas: Eureka______._. __ . ______._ .. __ 2.5(2.0-:3.0)/10 1.:3(J.2-1.5)/lG lJ(l1-lB)/ll ~ Oklahoma: 2.4(0-,1) /18 :Marble City_ ------.------_- _. _- __ . ______2.2/1 1.7 (1. G-1. 8)/G 11 (1 H2)/2 ______'I Stillwater______---.-.-.- .. -- - ______. _.. 2.5(2.1-2.8J/I,' LG(L3-L7)/I:3 12(11-14)/23 2.1 (1-3)/22 o Tcxas: Aransus. -. ------. --- _------_.. ' ______. 2.3(2.1-2.G)/211 1.6(1.(j-1.7)/2:~] 2(11-14)/13 2.11(1-4)130 ~ San Benito ______------.. -- .• --- ·------I1.G/l _~.------:3.5(3-4)/2 UJ Teague_____ ------__ .... ___ .. _____ ._ .. oO__ 2.G/1 1.7(1.7-1.8)/2 12/1 1.5(1-2)/2 Worthllln ______oO_____ 2.5(2.4-2.t3)/2 1.7(1.7-1.8)/3 12(11-12)/2 2.8(2-'lJ/G M . _,. __ .• ,.. I _....:.1 . ------_.- ___ __ >tI" >-: CUMC()IJ)I,S YARIIPJolNNIS ALBER'I'JolNSIS

Canada, Alberta: .. ------.. -.--...- --...... _-.. -_. -._- I "-~------.-'- ,..~------.- ~ ;> Lethbridge [241__ .' .... __ .... __ . ______.. __ .. ___ .. _.. __ .. __ .. _.. _: 2.4(2.0-2.6)/25 I I.G( I A-1.\1)/27 o Fort MacJ~eod [21______.. __ . ____ .. ____ .. ______]2(10-la)/51 ].7 (0-·1) /52 2.2(2.1-2.4)/2 Ui(1.5-1.G)/2 2 Brooks [1] _... ____ .. ______.. _.. _____ . __ .. _.. __ .. ____ . ____ _ 12(12-1al/:3 0.5(0-L)", () 2.3/1 ~, Montana: LuureL_____ .. ______.... _...,.. _.. J.G/ I 12/1 1.0(1)/2 South Dakota: OraL ______.. ___ ._ .... _._ 2.5/1 1.8/1 15/1 a.o/I ti 2.4(2. 2-2.ti)/.'i 1.5 (J.3-Ui) /6 12(11-1:3)/12 ,.:: Colorado: 1.2(0-3)/7 c:: Fort Collins__.. __ 2,4 (2. 2-2.7) /12 2 Fort Lupton_ 11.8(1.5-2.0)/15 12(lH:3)/15 L,1i(0-4) 2 I '-A 2.2(2.0-2.4}/3 1.8(1.(j-l.!)/2 1:3(1 L-IJ)/4 0.8(0-1)/5 • • • •

Nebraska: Lincoln______.. ___ .. __ • ______. 2.-1 (2.0-2. 8) /5 .I.8( 1.7-2.0)/7 13/·J 2.2(1-3)/5 .." Kansas: Stafford County ______. ______•. _. ______2.·1 (2.2-2.5) /·1 .I.8( 1.7-2.0)/.1 18(12-14)/{j 2.3(1-4)/4 p:: Oklahoma: Cleveland______. __. ______2.:3/t lAC 1.2-.I.G)/!i 10(10-11) /2 4.0/1 t'l Z -----. ...----- ...:...... ----- o CI·l.ll·OIIlE:'; YAllnl'~JNNISSONORENSIS ::tI ----- .." i i:I4 Mcxico: I Sonora, Aguu Calicllte______.. _ 2.2(2.1-2.:3)/10 1.2(1.1-1.-1)/8 11(1.0-1:3)/11 0.7(0-2)/Hi ~ Guerrero, Cocula ______. ______. 1.8/1 1.4(1.-1)/2 12(12-13)/2 1..5(1-2)/2 t'l l'uebla, l\fatamoros______. ______-.._ . ____ • ___ _ 2.1 (2.1)/2 1.4(1.3-IA)/2 12(12-1:3)/4 0.3(0-1)/3 § Nucvo Leon, Savin as Hidalgo______.. _.. ______2.:3/ I 1.5/1 13(1:3)/2 1/1 ;> Arizona: Z Brown Callyon______",_ . _j2.1(2.0-2.2)/+ 1.:3(1.3-1.'1)/2 14(I ..H5)/2 l.i>(1-2)/4 Vl Graham l\fountaill;i___ ... ______.______. ___ . ______.. 1.2/1 12/1 1.0/1 o Hot Springs______.. ______.. _ OJ ... 2.1 (1. 0-2.:3) /2 IA( 1.:3-1.5)/2 10(0- 11) /:3 0/2 Vl Patagonia _____ • ______. ___ • _.. __ ... __ ._ .• __ . 2.:3 (2.1-2.5) /2 1.'1(1.8-1.5)/2 ------..,.------_ ... "d __ __ t'l Sabino Canyon ______• . . _ _ _ . 2 . .1(2.1-2.2)/3 .1.2(1. H .2)/4 11(10-11)/:3 0/1 St. David ______.. ______. 1.4(0-2)/11 Q 2.1(1.0-2.5)/J2 IA(1.2-LG)/t7 12(U-l:3)/IH t'l Wellton______... _...... _ 2.1(2.0-2.8)/8 1.:3(1.0-1.3)/0 12(11-1a)/1G 1.l(0-2)/21 Vl Oak Creek______.. ____ . 2.-1(2.3-2.5) /a IA(1.3-L5)/15 la(12-14)/8 1.3(1-2)7 Parker ______• __ _ o 104/1 13/ t 1.0/1 "%j California: ·1------(') Indio______- .. _ . ___ .)2.:3/J 1.:3/1 12(12)/2 1.0(1)/2 d Shafter______. _____ . ______._ 2.2(2.1-2.2)/3 J2(10-l:3)/5 0.8(0-1)/7 Sangcr. ______." _' ___ . 2.2(2.0-2.-1)/1.0 J . .5(IA-1.8)/1G J2(11-14)/22 0.\)(0-2)/25 o~ Selma___ .. ______. 2.2(2.1-2.:3)/2 12(J 2)/3 1.0(1)/4 ""t::l Nevadu.: Rcno ____ ... ______. __ . _____ , 2.5/1 1.7/1 12( 12-1:3) /2 1.0/1 t'l "rashington: Vl Bucna ___- ______.. _- .. -- ______1 2.2 (2.0-2.:3)/3 .I .-1(1.·1-1.5)/:3 1:3(12-14)/5 ~ 0/5 ;> Lower Crab Crcek__ .. __ . ••. ____ • 2.2(2.0-2.:3)/2 1.:3(1.3)/2 12(J2-1:3)/2 O/J 'Moxee______2.1(1.8-2.-1)/1:3 ::tI ____ .. ___ 1.G(1.4-L8)/15 1:3(12-1.5)/23 0.8(0-3)/21 I=: Rocky Ford Crcek___ . __ . ______2.2(2.2)/2 1. 5 (1..5-1.7) /:3 1:3(12-1:3)/4 0/1 SlInnysic1c______2.0/1 1.7/1 J 2(12-1:3) /2 ~ Wiley ______... __ . ______-1 2.2(2.0-2.:3) / U 1.6(1..5-1.8)/12 J2(11-14)/10 0.8(0-3) /21 § I-l u:

t'>J 01 'I'AHl,g 3.-Measurements of lite subspecies oj the Culicoides nlriipcnnis cOinplerv (val'tle8 pl'(,8ented as "m.eon 1:\:1 (mini'llIJIl,ln-ma.1·imllln) /n'ltmber oj 1nea.s/lJ'ements") ~

('UJ.I(,OID~:SY.HtIIl'~:NNI;;SONOIl~:NSIS-Collt.illlled f-l M ('") . ; Third PIlIPI:;'-'T J,oclllit.y Teeth on AII t.eIII IlLI ~eg-mcllt j Wing length Illllndible sClIsoria ~ additional .... t" Utah: bi Lengthlwidth II j\fWjlllelt'r.~N'IImbm'.~ Delta __ .' NIt/IlIJ/!/' c:: NephL _____ . 2.5/1 1. g/I 1.0/1 t" 2.5/1 1.8/1 t"' Riehfil'lcL ~ ___ •• ___ .. 1.0/ L 1.8/1 13/1 ;.,t'l SOllth Salt Lake City_ 2.2/1 Utah Hot SpringR. 1.0/1 12(12)/2 0/1--- 2.511 2.0/1 14.(13-14)/2 9. Woods Cross. _ . 2.1 T'pxus: (1.0-2.2)/2 1.5(I.·H.U)/2 13(12-J.l)ja 2.0/1 -I Big Bend [30] .• 2.2(2.0-2.5)/07 Camp Stallley.• 1.2( 1.1-l.a)/Ga 12( I 0-15)/.10 n.8(0-2) /72 o Dryden ____ •.. 2.2( I.H-2.(j)/5 1..1(1..1)/6 12( I 0-13)/15 1.4(0-4)/12 2.1 (2.0-2.1)/-1 C; ]i'ort Da vis__ . .I.3( I.:H .'1)/:1 12(12)/3 1.5(1-2)/4 2.2(2.1-2.:3)/·1 Hovey •••• _. IA(I.:H.5)/4 12(11-I:l)/(j 0.4(0-1)/5 {P 2.1 (2.0-2.2)/:~I.a( 1.2-1 .. 1)/-1 .111110 •• ___ ••.. . .""------n.G(0-2)/7 2.1 (2.0-2.2)/2 .lA(.I.:H.4)/2 12/1 0.8(0-1)/4 o Kern'ille (L'!') _ 2.3(2.1-2.·I)/IG tr: .I..J-(1.2- I. 7)/8 12(0-1:~)/1.11.5(O-:l)f:H ](crn'ille Sla tiOlI 2.1/1 "d Palo Duro •• _ 1.'1/1 II (II) /2 1.0(1)/2 .." 2.·1/1 1.8/1 1:.~(la)/2 S:Uldl'r~on. _.• _ 1.4(1-2)/2 2.:~/1 1.·1/1 o Sonora Station •.•. , 1·1/1 I.O(O-:l)/2 'Tl 2.2(2.0-2.·1)/10 I A (1.3-I.U)/5 I·I( 12-15)/5 Weslaco.••. __ • __ ••• , 1.4(O-:l)/18 .... 2.5/1 1.2/1 2.0(2)/2 Oklahoma: Oklahoma Cit v •. 2.:l/l o Tenl1e,,~ee:.Pari~__ .__ . 1.7/1 I 1/1 1.5(1-2)/2 ::::l 2.2(2.2·2.:l)/2 ..., 1.2(1.1-I.;{)!2 I:l( 12-1·1)/:~1.0(0-2)/·1 a C1

~ tr: • • • •

I' 1,1('(111>1':l'\. ,\ HilI' I';:-;~IK 0['('11) 1;;\'1',\ 1.1l' (' 8 D:i ('aliforniH: t:'1 ~ Borax Lnk(' (20] 2.:1( 1.11 2.5).'28 I.(i( 1.0 I. ii!:l:{ I 1·1(11 I(j)/:li 0.5(0-:3)/oa 0 II 11£,11\,1111'2_, 2.2( 1.8-2.·1) '10 I.S( I.i I.II) !!l I:l( 12·1-1)/·1 1.0(0-2)/8 ::::..., \'('11 1.11rll 2 _ 2.2(2.2)'2 2. \(2.1 )/2 I:l( 12- I;{1I:{ 1.2( 1-2)f4 ...,T 131vI 11(' _ 2.1 (1.!l2.(i)/ii I.af 1.2 -I.:{,)·O I H( 12 " 1.1)!O 0..1 (0-1 )Iii Leil11a LindH 2.-1(2.·1 2.51/2 I.i ( I.i) 12 12( 12-1:11/2 1.5( 1-2l/" ;. ,...\;' Or(,lItl 2.2r2.0 2Ji) f I() !.il( 1.:1 I.Rl:!1 10(!l12)fii 0.5(0-2)/20 t:'1 It('~tillgHprillJ!;:< 2.2(I,!I 2.51,'2'1 1.0( 1.,1 l.i)112 1;)(11 15)/21 0.8(0-a)/2·' .... !:illI'll loga !:ipriIIg,; 2.2(2.0 2.512R 1.:\(1.2 1.5)':11 12(10 I:l)/:Hi O.i(O-2)/50 I') Baja California: ;. :Ells('uncin I.il I 1.0(1)/2 ~ Oregon: if, 'HIIIl1I1l('r Lak!' 2.2r2.0 2.:111 1.5(1 .•) I:HI2 1,1)/5 1.0( I) /., c: 1.')') 0:: Umatilla, 2.;{(2.2 2.,1}'2 12( II ·1 :1) ':1 1.0(()2) /4 if, F!'r11 Hidgp I.ii!l I "'I 1..5( I· 2)12 "'C \rat;hington: O'HlIllh'lI11 Dam t=; 2.0( 1.!l2.:{I;'!1 I.,j( I.-I I.il!1 1·1(12 15)'8 1.2(0,2)/10 :J

('allada, Briti~hColumhia: Kllmlo()p~ 2.2!2.0 2.1)!l I,M 1.:1 1.!ll,·I:l I:HI2 15)115 0.7(0 1)/2:1 t:'1 u; 0 2 Po~~ibl('di~tind 11l1l1-ili1lW popUlation of Illltil't('l'lIlilll'd I'II11k. "l (') r-; C (') 0 0 ~ Ul ;;:- :::: .... ;;; t:'1 ~ Y. if, -II>:'i 2S TECHXJCAL BlLLETIX I 170, r, 1', DEPT, OF AGHIC'FI.TT.'RE STATISTICAL EVALUATION!! OF FOUR CHARAC. TERS OF THE SUBSPECIES OF THE VARIIPENNIS COMPLEX • The tnxollomi~t JacC'd with thC' pl'Obl('1lI of "p]llll'atill~ a dose-knit taxonomic COJrlplC'x finds thC'I'(' al'C' llIany l)('r]1lpxin~ (]C'eisiolls to mak(', rBually 011C' 01' t\Yo ehal'nctC'I"; will 1101 bC' "u!lieient io sppanli(' all of' UIC' taxa PI'C'sC'n{ :-'0 that a 1l1l1ll1iC'r of charnel'PI's hn,\'l' I() 1)(' C'xalllinC'rl. find tho,.:C' <'11O:-:C'1I which di"cl'iminnte the bpst \)C'twr('n Ill(' taxa. thought to 1)(' pl'es('nL () f ('OUI'Sr, till' d('cisiolls nll'lIt iOlled nbo\'(' cnn hC', and ",-ual h' al'p. muclC' without, ill(' IlPlp of any f'tatisli('al lIl('thoclology, since the tax ollomist, often 11:1B no nwnn,.: of ('lIN'kill!! th(' a(,l'UI';I('Y of his clrcisioll:', l'x(,l'pt hy p:l'lH'l'al IlI'in('ipl('s, ('x]l('rirli('(', and intui'tion, This ,.:p('(ion iJlllsIl'a(C',s the 11\:11111('1' in which disC'I'iminnnt funC'fiOIl:' lIlay bl' llspd by till' t:tXOIIOlllisl' to elwck tllp a('('ura('\' of' his l'ubdi\'i sioilS, 'fllC' SI)p('i(':4 .is ('u{ir'oid('N 1'(I/'iiJi( 11 II iN (Coq,)' which is found tlll'ou!.!'hout tl1(' l'nilrd :::-;tntl'!", and in Canada lind .\[('xi('o, _\ ('li:':(,I'iminlmt fllll('(ioll is a lil1P:tl' flnH'tion in SP\'l'l':l1 "f1J'iahh.;.; that ",ill l'l'(ltH'p till' nll1ltiplp llH'aSIII'PIll('ni!" to a sing-Il' \'a111P, Tn the' I)J'p!:il'nt C','(:II II pIp, a di;;(,l'im i Ilam fUllction \\'a:-; ('a lc\llat'p(l for (':ICIt populatioll, Foul' qll:lntitllti\'(' ('1I:1l'a('II'I';'; "'PI'!' fO\lnd b." tlH' t ax olloJllists to han' potplltial \'alul' ",h(,11 IISP(] in ('onjuJl('rion 'witlt ('('1'_ lain qllnlitnti\'(, Clt:II':wtpI'S ill !"p]l:II'atillp: !!(,()!!I'aphieal population!". of tllP 1'lfl'iijlll1l1is t'olllplrx, Thl'sp t'llHl'f\('tl'I'S \\'('1'(': (1) X1,palpal J'atio·~~lell!!th of Sl'!!Illl'nl lH to its "'illfh, r~) X~, 1(,llgth of \\'ill!!, ill lllillillll'tl']'S, 1'1'0111 till' basal al'(,I""S to win.!! tip. (~) X, IJlllllbpl' of :llltl'llll:t1 !-('!,nlll'llts or tltt' !"PI'i('S I\"\'--\'J--\'Ir hrarin!! :"{'lIl'ol'ia, ' (I,) X 1,illJl1ll)(1I' of ((,pIll al tltl' apl'X of I Ill' Ill:IIHlihll', TIl(' iiI'S! IllP:tSlIl'PIlI('nts slIhl1lilfl'd l!\' till' taxollOlllisis \\'Pl'l' tak(,11 fl'OIll a ,,('rips of ~:3 spp('jm(>lls of I':ll'll oj' six 1)I'('sIIIllPd slIbsp(>('il's, Til OI'([PJ' f'o eOlllpll'tl' tIl(' ;'(,J'ips foJ' :-(,\'Pl'a \ :-lrI)"pl'('ip:, it, \\'as Ij('{'p"s:u',\' to in('llId(' spC'cinwll:-; f'I'OJll :-;catt!,!,pd hwaliti p ,> whi('h. as a !'psldt. of th(' (':tlclIlntiol1s. "'PI'P fOlll1d to I'('P 1'(':-('11 1 ,\I'P:I:- of illtl'I'gl'll(l:ltioJ1, To p:i\'l' tIl(' disl'I'illliJ1allt p(!llntioll!" :llld ,IH' l'p:-lJli;; 1'01' (11('!"e ('alclllation,; hpl'('. would 1>(' l'l']ll'titiol1s, and \Yollid S(,I'\'(, IH) IISl'flr! pUl'pO;'P sin('p thr)' \\'('1'(' lJa;-:('d Oil :til ('ITOI1('OIJI' l:txOJ1olllil' ]lI'Pllli:,:(', 'fill' I'rslilts of 111(':"r calelllalioJ1s \\'(,I'P \'C'I'Y POOl', with ('X('P;.;,;j\'(,\':ll'iatioJ1 indicated for ('\'ery sub;,pC'(,jps hilt "((l'iilill/I/is, ;\s a I'l'slIlt of this Work. til(' tnxonomists cOllelllsPIlIl'd ill table~, The sue('p~s of thp I'('-('\'aillatioll of 111(' "typicnl IO('nlit-ips" and ill(' •

• By .In(lson C McGuire, ;/1'" hiI)IlH?tl'il'iHIl, Ag-ri{'lIltlll'ul Hp1'(,:lI'('ll ~l'l'\'i('l', 'I'HE -,ORTJ! A:-'fEHJ('A~ :-;l'n:-;I'ECII';:-; OF (,I'LI('UI UES \'ARnpl~NNlS 20

:;;elect.ion of ,the "typien.1 f;(;,I'ip:;;" iB in for till' rOlli' 1l1PH"III'l'llll'lli!' :llhl 1'1(1('(' thai !-']l('ei men ill tlw sllbs]lPcips fOl' w\rich j 11(' \'alup of Lr i" tire la"l!est.

H,\' {'tdclIlnting th{, nrlw.'s of L t, for' ('Hch of thp l:>]l('{'illll'll;' il1\'oln'd ill tIll' \'Olll/lll!:al iOll of Illl' !\isl'l'illlin:tni rlllH'(ions tl)('III:,('ly('s. (hI' l'quntiolls al\()('aipd thp !'I)(·(·iII l{'Il;'. as shO\\'ll in talJlp ·1·. Thi;4 tnbll' jlllli{'tltps that t hl'l'L' an' fOlll' fairl.\· distill('( population;'. which ('all bl' ;:('pal'at('(l Oil tIll' IlH!'is of tIlt' foul' IIlpaSUn'III('lli!' 1I;,{'d ill tllt' allal~' si:-l. TllP filial S\'plu'atioll of (llis/I'ftih a 11\1 ((/I)( rtl /Isis IIIII:-:t lw InaIII' 011 t hI' ('ha I'act PI' () r tlil' lila 1(1 al'dl'a~,,!-'. ",hii'll is hal'\' ill IlIIs/ mlis alld !-'pinos(lill ((lb( r{f'IINio'l. •

TAIlI,I,; 4. IJll'clltla!JI of l1!J1cilllUIS IIs(d il/ 1111 alllll!J.~i,~ 1Il/0I'lI/utlo (([cit 1WbSll(·cils by 1111 rliscri/llilwit/ /Illtd/()I(.~

:\umh,f' of "p\'('i 111Ptl:-. /111 ,[I'lIli., u{.wl'lrll.

l'(lriipIIIJli., :~(i 07. ~ II () :.u; lIuxtrafts ~S () 100. () (J 0 .'Olwrell ~i" ~!I () () 11)(1. () () ,) 1111" rINl,~/,' -I :t 7 :-l\l, !I 0 7,1 (I('('irlc IItllhN :W (J :!(J, () II :-lO,O

In onll'l' \)(It/Pl.' to yi!'lI1i1izp (ll(' \'t'\atiollships :1I110 Il,!!' tIl(' li\-p :-uh !'peci{'s. I h,' di!'tull('P of ('ai'll sull!-'p('('ip,.; fl.'olll a II till' orlll'1' ,.;uiJ:'p('('i(':-< \\'n~ I'nl<>lil:1I('(1. and n (Iiag-ram (,Oll;,tl'll('({'(l. as :-:ho\\,11 ill ligHI'P II, t ou,hQli,

.l!'IGl:R1:: U.-- j){'g-ree of rellltil1I1ship~ of; fil'(~ :iubspel'if's of' till' 'l'(I"iiJ1(,lIlIi.~ ('oll1plpx, • as bused on 1Ill:1Iysis IJY diserilllin:lllt funelious. 30 TEC'JIXI('AL m'LLE'rr:-: 11;-0, t', S, DEP'l', OF M:BICl'LTFHS

.As fa I' as tile a Ilo('atioll of tilp Sp('('illl('ll!-' 1I!:'('(1 ill tht> :lnalysi:-; is ('011 ('PtllPd ancl the dish1!ll'PR tilr sllhspr('i(ls at'r from r:tt'h o(ilrt' in thp ding-rail!, the llnnly!:'is illdira!rs Ihat til(' rc-rn\lua(inll of thr ·'t<,"piral lo('nlitirs" \\"ns !:'1J('('p!:'sful: also tile' rp-!-'f'II'('(ion of til(' "fypi('nl series," • ~\Il()thrr t(>Rt whirh 1l1l1 \' hr nwdr is I hl' lise of discriminant fUI1t'tiolls on sp(>('in)(>ns frolll IOt'alities where inle'I'gl'adation is known ot' SllS !weted, This was dOIH' Tnt' Hi!l :'Pl'l'illle'll!:' of all liV(> ~Hlbspe('il's 1'1'0111 ~l (litl'HPnt !o('nlitie!:', Th(' l'Pslilts arc gin>n ill tablp :i, (;el1l'l'ally t11(> pOJllllat ions. \\'hpl1(>\'('r thl'l't' is a suflieil'llt .Illllllhl'I' or SPPCilllPIlS to Illakp IhI' P('J'('('llfagl's Illf>anillg'flli. fall wil'hill til(> sub;;preit's to \\'11 jeh till' taxllllOll1 i:,(s a 1I0('a II'(l thl'lll. Thl' f(l\\" ('X('PP lions that eallnot hI' pxpbilll'(l Oil olh!')' ('haradl'l':' 1l1;1,\' tltl'1l hp pla('('(\ tl littlP l110n tlcl'lIl'at('I.\' hy a:'slll1lillg IItnt 11](> :'ldlSI){'('il's ('xists ill a I'plati\'ply p:PIlt'til'all,\' PIII'(' !"tatf' nt till' .. t.",lien! locnlity." alld ('0\'('1';; an tll'PH ([('((,l'IlliIlPd Il\' ii~ (>(,()lo!,!'icHI 11(>(,11", .\;; a !il'IH'I'ul I'ltll' tltp fUl'th('l' (lIll' pl'l)('l'l'd~ n\\'tlY {'mIll I:ltp ('('nll'l' of dist I'illlilioll, til(> p:rt' of illt'(,I'g'mdatioll fl'OIll othl'I' sttl)SIll'l'il's \\'il II O\'PI'!appillP: dist:l'iiJut iOIlS, l ~~illp: til(' l)ouIHlal'.\' of till' I'Hng'l' of l'tlt'lt :-lIhsp('('it's n~ delhll>\[ in fip:lIl'l'l hy (Il(' laxollolllis(s. alld allo('atillp: prolmhilitil'S (IJa,;('t1 Oil ,I P(lis~()11 di;;ITihu(ioll. with Illl':ln ('(LU:!I to (JlW) alollg'l'adiallil1('s 1'1'011\ thl' ('('nl(,I':-; of !li:'t I'illllt i011. ('olliOIlI' linl';; (';111 1)(' dl'a"'" Oil a !li:'1 rihll liolllllal' ;;llO\\'ill,!.!' IIH' an'as \\'ithill whi('h l'Hl'h :'lliJsp('('i(':, 11I:1,\'\)(' (IX, pl'\'ft'd io (J('('1l1' ;w('o)'(lin,!i to !l,IL (),l~. !I,IHi, 'Illd IJ.(I~ limils of 1)1'01> ahility, ,'~ilh ;;lIdl n Illap. which of COIII'H' is lillli(pd in ifs, ;H'l'Ut'nc,\" IJ.\' (ht' :-:tlllpliIlP: ('ITor,,; of tIll' ('oll('('[iolls, thPlllsl''''P'':. il .is pOf'silJle to rpal\()('atl' "pl'l'illll'ns tls~ip:llpd to S\lhSI)('l'iP!" from Im'nlitip!-, f;u' 1'l'lllO\'('d t"I'OIlI thl'ir ('pniPI' of distl'ibution, Tltl'" i:-, ('~IH'('ially \'alllalJk ill till' ('n,,(' of ('lo:-'l'l.\' I'l'latl'd slIl>:-'\1\'('i(';. with IHlIHl\'PI'!:tppillP: distriilllti(lIls ;.IWIt tls ol'l,hl! II/(tf;" alHl (1I{"'/mli,,·({II)( del/Ris ill whi('h a l'OIl:-:idl'nthll' 11111111)('1' of :-'1 H'(·i III l'1l" a 1'(' !'olllld. 10 (j\'Pl'lap ill all 1'0111' ('\tal'actl'l':' ",illlul I:uIP()u-.:1 \-, By lI;in.!.!' sIIl'It pl'ohal>ility gl'adi('llb it i!" po!"silJll' 10 1'(I;ls;-:ign a Spt'('illll'll !"llvh its :\(), ;\ fl'OIl1 .\loXPl'. \\\I:-,It .. w\til'h Itas flH' following \';1 hiPS fOl' L, ; 1'1 I/'tlrii/II/lllis) 'co ~ I;,.!l' L~ (tlll.~t I'Iflis) '- ~(i~,S-I,

I J, (SOIlO,., /Isis) ~()f, f.1 1" (1111" rI('II,~ixl ~~Ii'-),I() L, I (/('('it/(/tllllh) ~Ii;),!)(i

I~.I' OUI.' 1'111(',.:, ill(' Sl)(>('illlt'll t"mlll which (II(' llll'aslIl't'IlIl'll1!" \\('I'P fakpll is If/tRim/is,

I L llls(>;ul of' usillp: L .. IJ.\' it:-,pl L till' quanf it,\' L r -j·lnI'1' is lIst'd. ",hpJ'l' ]>1' i,.: thl' pl'OfJnhility gin'll by I h!' ('Ol1iUIII'S. tin' following 1'('Sltlt" tLI'P OiJhlil1(,(\: 1,1 (,'adil/( IIlIix) ~1~.lll

L~ (/(/{.~t /'III is 1 ,c. ~!i,l.!);}

L (XI) /w /'(' /l,~ is J C ~(a, (Hi 1" (((/h( d(/I,~iN) "~~(jI.I!J r.,., (()('{'ithll/(tlis)~(i:i,(Hi • • •

TAB1J)~5.-··8u.bspeci('s allocation oj spccimells (d Ihe val'iil)(,7mis complcJ:jl'oln 21 loc(/'lities,inchulin{f those with >-3 ..... intel'{fl'(ula,/ion rJ., r-:: Z pprc~'ntllgcfnlling into- c ::q Xum!Jcr '? Subspecies Lot'aliiy ot ~Jl('<'i- ~ 1I1l'IIS t'aritjlClllli,q allslralisS(lIIOrellsi,~oCl"i£icntalis :> alberlensis I ,..,\;>' r-:: ~ :Jl!uSlb~rg,Tenl1- 10 (i0 ao 10 0 o uarUpclI II ill_ _ _ :> i lhl.ICT\'III<" Okla. fi no 40 0 0 Z I ___ I (l h.rlbournc, La 3 100 0 0 (fl 12 0 \12 0 8 C II IS 73 ! 0 l D tIl (llIslraU" __ •... '1"""'"''''I l~.ete~?b:lrg!Okl"Mo •.-- _ (fl i ;';,llb Iile, \ 11__ . ,,) 20 80 j 0 0 "0 Aransas, Tex._ .. _ 1(1 0 !It) 0 '10 r-: I o ,[ F~rlC:0ilinR; Colo. _ n 0 SO 0 711 ..., () t=J albcrlc/t.Si... .~_ ItOlal, H. Dak._ :3 67 33 0 (fl I HIllfford Co., Knm: -1 0 7fj 0 ' 25 () 'J-_il , 25 o ramI'""",I,y, '1'<', liO "l Kern-illc, Tex. __ II" 0 45 55 0 o SO/lorl'lIiis i St, David, Ariz._ a 0 0 100 0 Cj ) Wellton, Ariz ___ (l 0 II 100 0 t" ; Moxcc, Wash,3._ ... 13 0 ' 38 S 54 § 1 Wiley, Wash.3. __ .... . 10 (I 280 0 20 ,... (,YI""Cam., ____ .5 0 0 100 0 t:: _ H ucnerne,. <;Jalif___ . r-:: occidclI/a/iIL. 3 0 0 0 100 (fl j Orcutt, Cnllf.._ n 0 ' 8!.1 11 0 I Saratoga Spr., Calif.3." 21. 0 ' 14 86 0 ;; (l 217 83 :::l,.., I l{nll1loopR, H. C .. 0 0 ,... .~_._. ._L. __ ._.~_._,,_.._ . ~ _ 4<'_"_" __ ~_ "0 t':: 1 Subspecics Iwslralis and 1l/l)C"I('II.~iscan 1)(' st'pllr:t1ed onl,\' by lI1alt' characters. Z 2 Probably crrOIlCOUHand clln be ('orrt'cted by using probability contour" frolll distribution II1l1p. ....Z (fl 3 Assignment of "pecill1('n~re~i!iprilllnrily on charac(pr of IIlI~leIIcdeagus; pos~iblythesc specill1clls represent undescribed populntiolls,

CI.:I 32 'l'ECHt\lCAL BULLI!:'l'lN 1 t 70, t'. S. DI-:P'!,. 01~ AGIUCUUI'URE

These results place the specimen in orddcntalh. The re"ised break down for )[oxec, "rash., on th is basis is thereby:

l'en'PlllHI-(I! fallill!{ illto • XlIllIb('r of 8jwcillll'll» Imrt"ipcllllis li--:l:~l~:~~~- --- :~'~I:;e~:~;~---I-:('(~i::ali'; alberlcnsis I 13. - ~====~.=== -----~·l-----;-I-~------~fJ -1-----;-;

The coneeUon jH'ubabilities for .J[OXl'l', "~ash., hnsed on Illap contOlJl';-;, wouM be 0.7;1, for occidl'ntaZis, 0.0(; for 8011OI'('/18i8, and 0.02 for all othet· subspeciN;. -,\.llything oui:Bi

Ecological Variation: The individuals of 1Jariipellnis f". str. hom the cooler northeastern States are eonspicuonsly l:u'ge, with wing length avcraging 1.7 111m.: albm'tl'7!sis from t11e northern prairies is next largest, with wing length of 1.(i mrn.: California populations of oceidentalis nearest the cool Paci fie Coast are of Janrer size than those OCCUlTing in the Wimner interior, :mel British Columbill and "'ashi ngtoll indi \'iduals are a]so larger. Tlw smaJlest ind ividua,}s of IIlis comph'x arc found ill populations of sonm'ell8is from the very hot desert areas of A rizona and )wighbol'ing States (wing length 1.2 mm.), while popUlations in the northem pad of the range of this subspecies h:we wings ayeraging lip to 1..() mm.long, notably at 'Vi ley, 'Yasll. ,Ye ]Hlve alrClldy pointed out that specimens of 801101'(?IlRis at Kel'lTille, Tex., taken in winter and in the eool, early-sprillg season are larger than those taken latel' in the summer. The snbspeci('s of t'Cl1'iipenni8 ttpparently have dew>loped in ac('ol'llance with Bergmann's ru](' (Dobzhansky, 1$)41) tltnt races in cooler climates are larger in body

than those of the same spl~cies in W:U'mel< climates. ~ w In the variipennis eomplex, the darkest individuals are found )n the subspecies Nll'iipennis of the humid forest rcgions of the eastern and northerll part of the country. Variation in color genemlly pro ceeds in a gradient tlu~ough populations of the various subspecies from east to west and north to south, until in the extreme ariel South west the dark wing markings become yel'y l'ecessive in sOlwl'ensis and occidentalis. However, there are quite a number of specimens from • coastal localities in California, doubtfully assigned to occidentalis, 'l'HE NORTH AMERICAN SUBSPECrES OF CULIComES VARllPE1I."'NIS 33

which are lar~e and very dark. 'l'hese come from the cool, humid, maritime localIties, such as Hueneme, Ventura, Novato, and "Theeler's Springs. This geographical pattern apparently fits Gloger's rule (Dobzhansky, 1941) which states that in insects racial variation in • pigmentation increases in humid and cool climates and decreases in dry and hot climates, the humidity apparently being more effective than temperature. Phylogeny: We would unhesitatingly split the 1Ja1'iipennis ('om plexiirstinto two main stocks: the subsp~eies rCll'iipennis of the north eastern forests, and s011.m'ensis of the southw~stern urid country. The subspecies variiZJenni8 has undoubtedlY existed for a long period 01' t.ime in a. relatively stable and equable environment, as indlca.ted by its relatiye lUliformity of characters. On the other hand, the sonOl'en Ris stock has deyelop~cl, or retained, a wide range of genotypes, pre sumably adapted to fluctuations in the many isolated and limited en ,rjronments of the Sonthwest. In the periphery of the range of 801101'ensi,, some of these ada.ptive t.YP~s have evolved into the closely related subspecies (LustmZis, albel'tellsi8, and occidentaNs, as weLI as many other Jesser populaJions, which we cannot yet recognize with our limited matel'ial. Becnuse the range of characters is greater, and the intergradation is much more gradual, it appears to us that in the zon~s of geographical contact t'he genetic bal',riers between t.h.ese del'lyed forms and 80nOl'('1!,sis fire nnwh l('ss than those scparatll1g them from 'l'(ll'iipennis. Thus, "'c bclie"e thnt the three major pedph ('ral populations Wllich we haw wnturcc1 to name as subspecies 'were dill'el'cntiated from sonorel1Ris stoek aftcl' the subsperies I'a?'iipennis hacl renchccll'clatiye stability. Problems: As this study in'ogressed it became apparent that thel'c did exist some geographical and ecologieul co]']'('lation with certain characters. 'Ye 'were faced with the prolJlem of obtaining nc1cqnate: series of specimens to determine trends of vttriation and to define as closely as possible the zones where cliscontinuiti~s appeared. Requests 'for additionalmatedal on short notice bronght remarkably good re "ponse considering the general sctU'city of Culicoides in Departmental collections. IIoweyer, there stilll'emain many areas from which poor material or inadequate series has seriously 11inclered our study, a nel wl'ak~netl Olll' conclusions. n should be acl(h'd that ill zones ol'in tl'l'gl'adatioll light-trap maicri:ll is usually illadl'quatp IInl~"s it ('an 1)1' a"ccl't::1i I1PCl how ilia ny ty}>l'S of bl'(,Ptl ing si tun.tion a 1'1' bl'i np: sa 111 piNt '1'l1e primal',\' :l.l'l'tl" wh ieh J1l'ed intellsiyc study inelud~ : 1. The Xe\\' England area, ",hel'e the complde abspn('p of l'N'ol'\I:: j)l'(wents OUI' (Jp(ining tlH' nOI'l'lwn:-:(p\'ll limit of tllp I'ange of /'{()'iip('I1 !lis s, str. :2. Thc Southeast', l'slw('in Ily the Ca 1'01 ina" a nll Florida. ,,'hel'(> till' (~xt.I'Cll1e :-;oul"hel'n 1i Il1 it of dist:t'ibnt'ion of the ('ompl~x is lInknown. and wlll'l'l' tlw infl'I'I'('1af'ionships of th~ snbspl'C'ies 1YlI'iiju>Jll1is. I(IlS tl'Cl.tiR, and S()IIO/'(,IlSis nlust he wOl'ked olllin detail. A]l illl])nl'i'nnt question here is whefhl'r 01' not thp main ditt'('\'el1('e is ecol()~i('al. p. !r.• saline enyirOnlllent. ' , :1. The southel'n (:r\'~af' Plain" (Oklahoma-Kansas HI'en) whpn' fht' I'elation b~t\\'een (w.~ti'alis and ("7J(,I'tI'1I8i.~ is ohs('url', ·k The nOl'thl'I'n prail'ies, whel't' the natt!!·p of tl1(' lilH'. 01' al'ctl. bl' • twcen the rangps of alb('!'IeI18r~ nnd '/'{U'iiprnnis s. stl', is unknown. Is 34 'l'ECHKICAI" B"C"LLETlX 11 ,OJ Co S. DEPT. OF AGRrCl'LTURE

it as simple as an ecological boundal")' between forest n!ld prai rie t lYe ha,e yjrtually no collections from this cri lica I area rx(end j ng- frOI11 Saskatchewan and :Minnesota south to Kansas tlncl Iowa. 5. The Rocky )Jounbtins. Does l'w'iipCllnis s. str., as we sllspect', • 1't1l1ge down the Rockies at higher elerations into ('olorado? .\gain) 110 substantiall'ecords are anlilable, 6. The Great Basin, "That is the lIatllre of the apparent difl'eren iiation of sonol'el1sis in l~tah into R populatioll with larger \\'ing sil':(' and possibly other eha I'llrfel's { 'I'he enti 1'1' J)OI'fIt('I'n (hpa I' Basin must be samplNl s},stpll1atirally for a poplllation anal.n;is :111(1 l)iolog-ieal study. i, 'Tlm Pm·jli(, Coast, ,rhat is tIl(' ('('ologi('nl sppal'afioll behrl'('11 ·~()]I0]'(']Mis and oreir/ellfalis ill tIl(' l'a('ific Coast Statl's, and wltat lilnits the p,nstel'n disiribntion of orf'idclltltlis? R. Ca Ii1'01'11 ia, Is tlwrl' a disti Ilrt ('oast a.1 poplll:ttion 0 f t'hc HIll'IIPIlII' type that ('nn hp spparat(·(! ('C'ologi('nlly fl'oill tYl'i('al orridell/rili8! n, XorthprJ1 ('Hllada, "'hat is tlIP Ilol'thpl'll lillli(" of distribution or this complpx! "'Ire hnn too few Canadian ('01 l('cf ion.s to e\'en ypn/uf'(' a guess, 10. 1[l.'xieo and Ct'nt 1';) I _\llIf'ri(,:I, ",,'hat is flIp souiht'1'1l 1i111 it of distribution of tllis ('ompkx, IlO\\' 1'('('ol'dl'(1 :IS far SOllt h :IS tllf' ~\rl'xi('an Statp of GIlPl'l'('I'()! Dol'S onl)' (]('('ilhll/rdi.v 0('('111' in Baja California! .\rp then' fUI't:hp)' sllrnll. isol:li('d subslwei('s ill tlj(> )/(oxi('all hi;:!'ldands, as in other gToupsoT animals! TJwse Hrp all impol'hlllt qups(iolls on n'hi('h 'n' ('a II 1101' (,,'P11 "Pllt II1'P to !!IIPSS an answPl' wilhollt I'xtenSiYl' lipId work. 0111' sfud,- ll:ls so -rar I)(,PI1 I'Pstl'i('t(>{l yil'lil:dh' to that of a IIIUSPIIIII taX()IHllllist. ','"p \)plip\'l' that:, we ha \'(> 1'P:1('1Ip(l ni(, I))'adil':d Iilll its ()ft hnt Ii Ill' of atiapk hyitspH, :mcl flll",hl'1' nnalrsismnst I>p illfimat'pl" t'i('([ ill with clptailpcl biolo![ieal stnclips, Tllp e(,~l\l()lllic jllstili('ntioll f~lI' slwll Sind." pl'obably J'psts 011 the OUleOlllp ofprojP('('(] stndi",; on thp tl'anSI])issioll of :tnilllni cliseasps by this common and widespread SIW\'il'~ . .sul'h ns ill(' CUlTent W()I'I~ 011 hlJ]('tongm' of Rhppp. 1rO\\'("'PI', flIP alJlm<1a n('p :l ))(1 IIl>iqll it)- of th(' I'lIl'iipflll1is ('omplex .ill nahll't'.ils adaptnlin)) to (lin]'s!,' biotic ('ondi tio))s, and tllp appnl'('nt taxollOl1li(' (litJ'(>[,Plltiarion in pnl'alkl to sudl ada pta lions. togpt:iWl' wi III its l'pla t iy\' \'as(' of ('llliul'pin thI' labol'ntol')', ]Joint. io Ihis ('()lJ)ph'x as OJ1(> of thl' I!lOst Pl'lllllisillg' RlIbjpds :/'01' tlIe Rtud~' 01' r\'(llutiOtl in the [ield of 0ntolllolo!!y. REFERENCES TlM'OX. :'If. J!);j;t A S'ITny 01' 'l'HF. AII1'HIIO\'()IIS O~' :lmmCAT, ,\X!) YETElIlX,\HY nIPOHTAXC1:: J:i ,'In; COlX:I( /!L\ BASTX, "\'flsh, Agl'. Bxpt. fHa" ~r('('h. Bill. 11, ·10 pp. BELlO X • ,T, :s'" AX\) :'ITC'nOXAL!), "Yn.r.r.\)l A, 1f);jU. ,\ I'OI'I'I,,\Tl()X Ill' l'II.\XO'I'.\I';Xl.\ ,IXllYIiOlt FI:O~1 111':.\'1'[[ Y,\Ler;Y. WITH ngSC1UPTlOXS OF ,\LL ST.IG\'S ,I;;n [nSCt'ssmx 01" Tln; C'O:lrpr.r.x (UIP'I'EI:A, Cr'LH'Ill,\r.l, l~lIt. ~O(', ,\ll)CI', .\nn, 4n: 103-18:!, TJOB7.H,IXSKY, TH, n. 1\),11, ()EXETICR .\;;ll THE OIl[(;IX OF SPE(,T!::S. ·J,Hi P])., illns. E(l. 2, rcl'. ('olulIIhin r-nh'PI'Rit),l'I'Pss, XI'\\, York, --.\XlJ EI'LlXG, ('. 1n-l,l. COX1'IIH1UTlOXS ~·O THE m;N~;'l'I(,S, 'I',\XOXO~IY, ANIl 1,('or,OGY 01'- rmoso, l'rrILA PSI,eIlOOBS('{'ICA .IX\) ITS RELATJrES. Carncgic lnst, ""1\1;11, rub. fJ54, 183 llP, • 'rUE XOHT.H, A~II~HIl'AX ~UI3P,PI'l'IES OF cn,ICOIDES \-ARUPENNlS :35

DOWSES, ,T, A, 10;)0, ,IIAIIITS ASJ) 1,li'I';-('Y('U: OF ~TI.I(,OIlJI':S Sl'In:(TL08l'8 (~IG,) XatllL'f', London 1(;(;: ;:; I 0-(;11 .

H););), OIlS~:ll\'A'I'lOSS os TJII~ 8\\,,\1011.'$: 1.'I,IC;II1' .\SI) ~L\'I'lSG O~' el'I,ICOlll~;S tIHPn:IIA: (,EIIAT()I'OGO:"l'lIMI~), Hoy, Enl, So('" London Trans, IOU: • :!1a-23G, ~ pIs, EIlW.\lWS, ]", '\"" Or,UIlOYIJ, '1 L, ,\XII S~L\ll'I', ,l. HI3!), 11111'1'1$11 BI.nOIJ-HI'('KIXn l'LIES. Bl'iL~llIs. (Xnt.llisL, U(·pt. Zoology), J:i!i pp., ,I;) pIs. London. 1·'n;uLEH. n. (;. II. HI:-,1. TIII';S()('TII.\FIIJ('.\:\ IIITIS'; \I II",,·;s OFTIII';lil,:sn; ('('LJ('OIlI~;S '(,~:H.\T()- 1'()(H1SII)., J)II'T,), ()ncll'l's(ppOnl'l ,/0111'. \'(,1'. nt';:. :!;) !:!) : ::. ;{a. 1\\,.\'1'.\, ~I, tn:{;;. os III.OOJ)-Sf TI":\"IO IIII'TI':HlIn; I:" SI';I''I'S, II it'i1"h illl:l-Kol\('h II-J)okolw i K:li;:hi:!: ;;O!I, 1111 ,1:1 P:I 11('$1'.] KII.\L\I-'. K. 'I'. tn:i:!, TIll'; ('1'1,(('01111';8 OF 'rilE WI('IIIT.\ 11I':Fl"i!':. "h:I.,\IIi1\1.\. T'\X():"')~IY .\:"11 :"E.\snS.\I. 1:\"('II)(o;:"l'\o; IJ)II''I'EIII.\. IIEI.~;I$."':I. 1':11(. Hol', .\111('1'. _\1111, I;;: :W,·;{:iK I'llWE, I), A .. ,,:"J) I 1.\11IlY , \Y_ 'I'. 'I!);;,!. ISOI_\'I'IO:"I' (II<' '1'111'; lII.n:'ln,t;t'I'; ""U'S Jo'I{t)\1 TI';X.\8 SIII';~;I" ··('n,((·OIll~$ s II ow:\" '1'0 III;; A YE('TOII. .\ IlIl'I·. \'pl. ~I pd.•\~SIJ(' ••loul'. 'I :!-I : ~:i:i-,:!:iS. HOOT. I". '\1., .\XlIlIoFI'~I,\X. \\'••\. 'W;ri. Tla: XOIITII .U"':HH'.\ ~ Sl'\o:l'l ES OF ('( LIt'UIIlEb. .\ lill'l'. ;'0111', lin!'. :!:;: 1;;0 I,(j. S pIs. 'I'nKl';\',\(i.\. ~1. 'W;),. ::;,\SII Fr,n:!; .('EIt,\·I'OI'Ol;(I;\'III.\Jo;, IIII'TEIU, FIlO\1 .I.\I'.\;\,. Tl'IlUII'l'do 1: :!;{;~':1:{'. Ii pIs.. :1:{ Ii)!s. "'.11ln:III':.ul. F. I·~. 'J!);H. 1I,\.\I.\(;t; TO l.ln;STIH'K Ill' 111.0(111->;('<.'''1:\"(; .\IIll(iI':S. Okla.•\p:. l~xJlt. Xla. HpL.W:t! ;11. JlJl. :!Ii-I-:!OS, \\-mTIr, '\"- W. lfl;;2. TH~: 111';1,1':111.\1'; OF (',\I.IFIIW'\I.\. (':lIil'. "IIiI'. 1'II11s.. gilL n: n:;-:!(;ii. -- .\:"1' II BOT'I'DI "It, L. ,I. l!l;)G. .\ 1'()I't' ( ..\TIOS S'ITIlY OF TIll'; IT I.Il'OIIlE!; \( 1Il(;t·;s 1Il.lI''I'I';H.\, 11 "1.~:lIJ.H:1 OF TII~: 1':Il\L\lWSI'L.\'I'E.\l: 1l~;(ilO;\, OF 'I'I':X,\S• .\IOS'IlIii'1l Xl'WS 1U 1,,11: :!;;!i·:!un. 10 ligs

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